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GENES ARE SOLAR ENERGY IS BOTH

WITHOUT LIMIT AND
SELFISH WITHOUT COST
MOLECULES
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ON A DIFFERENT TO BUSY ITSELF WITH
TIMESCALE THE EARTH ITSELF
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BURNING
ECOLOGY
QUESTION
ALL BODILY
ACTIVITY DEPENDS
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BOOK
BIG IDEAS SIMPLY EXPLAINED
THINK GLOBALLY,
ACT LOCALLY WE ARE
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THOUGH
WE ARE PLAYING DICE WE HAD
WITH THE NATURAL ANOTHER
IF YOU DO NOT KNOW THE NAMES OF ONE TO GO
ENVIRONMENT THINGS, THE KNOWLEDGE OF THEM TO
IS LOST
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FOREWORD BY
TONY JUNIPER
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CONTRIBUTORS
JULIA SCHROEDER, CONSULTANT DEREK HARVEY
Julia Schroeder received her Ph.D. in Animal Ecology from the A naturalist and teacher with a particular interest in evolutionary
University of Groningen in the Netherlands. From 2012 to 2017, biology, Derek Harvey graduated in Zoology from Liverpool University
she headed a research group at the Max Planck Institute for in the UK. He has taught a generation of biologists and led student
Ornithology in Germany, studying social behavioral ecology. Julia expeditions to Costa Rica, Madagascar, and Australasia. Derek now
currently researches and teaches evolutionary biology at Imperial concentrates on writing and consulting for science and natural
College London. history books.
CELIA COYNE TOM JACKSON

Celia Coyne is a freelance writer and editor living in Christchurch, A writer for 25 years, Tom Jackson is the author of about 200 nonfiction
New Zealand. She is the author of Earth’s Riches and The Power of books for adults and children and has contributed to many more. Tom
Plants and writes and edits articles on science and natural history studied zoology at Bristol University, UK, and worked in zoos and as a
for magazines, newspapers, journals, websites, and books in the UK, conservationist before turning to writing about natural history and all
Australia, and New Zealand. Her aim is to make scientific subjects things scientific.
accessible to lay readers.
ALISON SINGER
JOHN FARNDON
Alison Singer is a Ph.D. candidate in Community Sustainability at
The author of hundreds of books on science and nature for both Michigan State University, US, where she studies storytelling and
children and adults, John Farndon studied geography at Cambridge science communication. She has a broad educational background in
University. He has written extensively on earth sciences and the writing, ecology, and the social sciences. Alison has worked as an
environment, focusing in particular on conservation and ecology. educator for environmental charities, and for the US Environmental
His books include The Oceans Atlas, The Wildlife Atlas, How the Earth Protection Agency.
Works, and The Practical Encyclopedia of Rocks and Minerals.
TIM HARRIS
After studying Norwegian glaciers in college, Tim Harris traveled the
world in search of unusual wildlife and extraordinary landscapes. He
has explored the dunes of the Namib Desert, climbed Popocatépetl in
central Mexico, camped in the Sumatran rain forest, and searched
the frozen Sea of Okhotsk in Russia. He is a former Deputy Editor of
Birdwatch magazine in the UK and has written books about nature for
adults and children.
CONTENTS
12 INTRODUCTION 34 We’ve discovered
the secret of life
The role of DNA
THE STORY
OF EVOLUTION 38 Genes are selfish
molecules
The selfish gene
20 Time is insignificant
and never a difficulty
for nature
Early theories of evolution ECOLOGICAL
22 A world previous to ours,
PROCESSES
destroyed by catastrophe 66 The fitness of a foraging
Extinction and change 44 Lessons from animal depends on
mathematical theory its efficiency
23 No vestige of a beginning on the struggle Optimal foraging theory
—no prospect of an end for life
Uniformitarianism Predator–prey equations 68 Parasites and pathogens
control populations
24 The struggle for existence 50 Existence is determined like predators
Evolution by natural selection by a slender thread Ecological epidemiology
of circumstances
32 Human beings are Ecological niches 72 Why don’t penguins’
ultimately nothing feet freeze?
but carriers for genes 52 Complete competitors Ecophysiology
The rules of heredity cannot coexist
Competitive exclusion 74 All life is chemical
principle Ecological stoichiometry
54 Poor field experiments 76 Fear itself is powerful

can be worse than Nonconsumptive effects
useless of predators on their prey
Field experiments
56 More nectar means

more ants and more ORDERING THE
ants mean more nectar
Mutualisms
NATURAL WORLD
60 Whelks are like 82 In all things of nature
little wolves in there is something of
slow motion the marvelous
Keystone species Classification of living things
84 By the help of 138 Life is supported by a vast

microscopes nothing network of processes
escapes our inquiry Energy flow through
The microbiological ecosystems
environment
140 The world is green
86 If you do not know Trophic cascades
the names of things, the
knowledge of them is lost 144 Islands are ecological
A system for identifying all systems
nature’s organisms Island biogeography
88 “Reproductively isolated” 150 It is the constancy of

are the key words numbers that matters
Biological species concept 114 Birds lay the number Ecological resilience
of eggs that produce
90 Organisms clearly the optimum number 152 Populations are subjected
cluster into several of offspring to unpredictable forces
primary kingdoms Clutch control The neutral theory of
A modern view of diversity biodiversity
116 The bond with a true dog
92 Save the biosphere and is as lasting as the ties of 153 Only a community
you may save the world this earth can ever be of researchers has a
Human activity Animal behavior chance of revealing
and biodiversity the complex whole
118 Redefine “tool”, redefine Big ecology
96 We are in the opening “man”, or accept
phase of a mass chimpanzees as humans 154 The best strategy
extinction Using animal models to depends on what
Biodiversity hotspots understand human behavior others are doing
Evolutionarily stable state
126 All bodily activity
THE VARIETY depends on temperature

Thermoregulation in insects
OF LIFE
102 It is the microbes that ECOSYSTEMS
will have the last word
Microbiology 132 Every distinct part of
nature’s works is
104 Certain tree species necessary for the
have a symbiosis support of the rest
with fungi The food chain
The ubiquity of mycorrhizae
134 All organisms are
106 Food is the potential sources of food
burning question for other organisms
Animal ecology The ecosystem
156 Species maintain the 170 I have great faith

functioning and stability in a seed
of ecosystems Ecological succession
Biodiversity and ecosystem
function 172 The community
arises, grows,
matures, and dies
ORGANISMS IN Climax community
A CHANGING 174 An association

is not an organism
ENVIRONMENT but a coincidence
Open community theory
162 The philosophical study
of nature connects the 176 A group of species THE LIVING EARTH
present with the past that exploit their
The distribution of species environment in 198 The glacier was God’s
over space and time a similar way great plow
The ecological guild Ancient ice ages
164 The virtual increase of the
population is limited by 178 The citizen 200 There is nothing
the fertility of the country network depends on the map to mark
The Verhulst equation on volunteers the boundary line
Citizen science Biogeography
166 The first requisite is
a thorough knowledge 184 Population dynamics 202 Global warming isn’t a
of the natural order become chaotic prediction. It is happening
Organisms and their when the rate of Global warming
environment reproduction soars
Chaotic population change 204 Living matter is the most
167 Plants live on a different powerful geological force
timescale 185 To visualize the big The biosphere
The foundations of picture, take a
plant ecology distant view 206 The system of nature
Macroecology Biomes
168 The causes of differences
among plants 186 A population 210 We take nature’s services
Climate and vegetation of populations for granted because we
Metapopulations don’t pay for them
A holistic view of Earth
188 Organisms change
and construct the 212 Plate tectonics is not all
world in which havoc and destruction
they live Moving continents
Niche construction and evolution
190 Local communities that 214 Life changes Earth

exchange colonists to its own purposes
Metacommunities The Gaia hypothesis
218 65 million years ago 270 The introduction

something killed half of a few rabbits could
of all the life on do little harm
the Earth Invasive species
Mass extinctions
274 As temperatures increase,
224 Burning all fuel the delicately balanced
reserves will initiate system falls into disarray
the runaway greenhouse Spring creep
Environmental
feedback loops 280 One of the main threats
to biodiversity is
infectious diseases
THE HUMAN 242 The chemical barrage
Amphibian viruses
FACTOR has been hurled against

the fabric of life
281 Imagine trying to build
a house while someone
The legacy of pesticides keeps stealing your bricks
230 Environmental pollution Ocean acidification
is an incurable disease 248 A long journey
Pollution from discovery 282 The environmental
to political action damage of urban sprawl
236 God cannot save these Acid rain cannot be ignored
trees from fools Urban sprawl
Endangered habitats 250 A finite world can support
only a finite population 284 Our oceans are turning
240 We are seeing the Overpopulation into a plastic soup
beginnings of a rapidly A plastic wasteland
changing planet 252 Dark skies are now
The Keeling Curve blotted out 286 Water is a public trust
Light pollution and a human right
The water crisis
254 I am fighting
for humanity
Deforestation
260 The hole in the ozone

layer is a kind of
skywriting
Ozone depletion
262 We needed a mandate

for change
Depletion of natural resources
266 Bigger and bigger boats

chasing smaller and
fewer fish
Overfishing
306 The time has come for 324 We are playing dice with
science to busy itself the natural environment
with the Earth itself The economic impact of
Environmental ethics climate change
308 Think globally, 326 Monocultures and

act locally monopolies are destroying
The Green Movement the harvest of seed
Seed diversity
310 The consequences of
today’s actions on 328 Natural ecosystems and
tomorrow’s world their species help sustain
Man and the Biosphere and fulfill human life
Programme Ecosystem services
312 Predicting a population’s 330 We are living on this

size and its chances planet as though we have
of extinction another one to go to
Population viability analysis Waste disposal
ENVIRONMENTALISM 316 Climate change is
AND CONSERVATION happening here. It
is happening now 332 DIRECTORY
Halting climate change
296 The dominion of man
over nature rests only 322 The capacity to sustain 340 GLOSSARY
on knowledge the world’s population
Humankind’s dominance
over nature
Sustainable biosphere
initiative
344 INDEX
297 Nature is a 351 QUOTE ATTRIBUTIONS
great economist
The peaceful coexistence
of humankind and nature 352 ACKNOWLEDGMENTS
298 In wildness is the
preservation of the world
Romanticism, conservation,
and ecology
299 Man everywhere is

a disturbing agent
Human devastation
of Earth
300 Solar energy is

both without limit
and without cost
Renewable energy
FOREWORD
As a small child, I was fascinated by nature—birds, I am delighted that Dorling Kindersley decided to produce
butterflies, plants, reptiles, fossils, rivers, weather, and much The Ecology Book, setting out the key concepts that have
else. My youthful passions set me on the path to being a helped shape our understanding of how Earth’s incredible
life-long naturalist, and to working as an environmentalist, natural systems function. In the pages that follow readers
studying the natural world and promoting action for its will also discover something about the history of ecological
conservation. I have worked as a field ornithologist, writer, concepts, the leading thinkers, and the different perspectives
campaigner, policy advocate, and environmental advisor. All from which they approached the questions they sought
of these diverse interests and activities have, however, been to answer.
linked by a single theme: ecology. One thing that sets this book apart is the manner
Ecology is a vast subject, embracing the many disciplines in which the rich, memorable, and attractive content
needed to understand the relationships that exist between is presented. A huge body of information and insight is
different living things, and the physical worlds of air, water, effectively conveyed by clear layout, graphics, illustrations,
and rock within which they are embedded. From the study and quotes, enabling readers to quickly achieve an
of soil microorganisms to the role of pollinators, and from understanding of many important ecological ideas and
research into the water cycle to investigating Earth’s climate the people behind them: James Lovelock’s Gaia Hypothesis,
system, ecology involves many specialist areas. It also Norman Myers’s warnings about impending mass extinction,
unites many strands of science, including zoology, botany, and Rachel Carson’s work to expose the effects of toxic
mathematics, chemistry, and physics, as well as some pesticides among them.
aspects of social science—especially economics—while The diverse body of information found in the pages that
at the same time raising profound philosophical and follow could not be more important. For while the headlines
ethical questions. and popular debate suggest it is politics, technology, and
Because of the fundamental ways in which the human economics that are the vital forces shaping our common
world depends on healthy natural systems, some of the most future, it is in the end ecology that is the most important
important political issues of our age are ecological ones. They context determining societies’ prospects, and indeed the
include climate change, the effects of ecosystem damage, future of civilization itself.
the disappearance of wildlife, and the depletion of resources, I hope you find The Ecology Book to be an enlightening
including fish stocks, freshwater, and soils. All these overview of what is not only the most important subject, but
ecological changes have implications for people and are also the most interesting.
increasingly pressing.
Considering the huge importance of ecology for our
modern world, and the many threads of thought and ideas Tony Juniper CBE
that must be woven to gain an understanding of the subject, Environmentalist
INTRODU
CTION
14 INTRODUCTION
F
or the earliest humans, Swedish botanist Carl Linnaeus organisms change over time, and
a rudimentary knowledge developed a classification system, even become extinct. The
of ecology—how organisms Systema Naturae, the first scientific Frenchman Jean-Baptiste Lamarck
relate to one another—was a matter attempt to name species and proposed the first cohesive theory
of life and death. Without having group them according to of evolution—the transmutation
a basic understanding of why relatedness. Throughout this of species by the inheritance of
animals grazed in a certain place time, essentialism—the idea that acquired characteristics—in 1809.
and fruit-bearing plants grew in each species had unalterable However, some 50 years later it was
another, our ancestors would not characteristics—continued to Charles Darwin—influenced by his
have survived and evolved. dominate Western thought. experiences on the epic expedition
How living animals and plants of HMS Beagle—and Alfred Russel
interact with each other, and Great breakthroughs Wallace, who developed the concept
with the nonliving environment Geological discoveries in the late of evolution by means of natural
interested the ancient Greeks. 17th and early 18th centuries began selection, the theory that organisms
In the 4th century BCE, Aristotle to challenge the idea of essentialism. evolve over the course of generations
and his student Theophrastus Geologists noted that some fossil to adapt better to their environment.
developed theories of animal species suddenly disappeared Darwin and Wallace did not
metabolism and heat regulation, from the geological record and were understand the mechanism by
dissected birds’ eggs to discover replaced by others, suggesting that which this happened, but Gregor
how they grew, and described Mendel’s experiments on peas
an 11-level “ladder of life,” the first pointed at the role of hereditary
attempt at classifying organisms. factors later known as genes,
Aristotle also explained how some representing another giant leap in
animals consume others—the first evolutionary theory.
description of a food chain.
In the Middle Ages (476–1500), There are some 4 million Making connections
the Catholic Church discouraged different kinds of animals and The relationships between
new scientific thought, and human plants in the world. Four organisms and their environment,
understanding of ecology advanced million different solutions to and between species, dominated
very slowly. By the 16th century, the problems of staying alive. ecological study in the early
however, maritime exploration, David Attenborough 20th century. The concepts of
coupled with great technological food chains and food webs (who
advances, such as the invention eats what in a particular habitat)
of the microscope, led to the and ecological niches (the role an
discovery of amazing life forms and organism has in its environment)
a thirst for knowledge about them. developed, and in 1935, Arthur
INTRODUCTION 15
Tansley introduced the concept New concerns botany, and their microdisciplines,
of the ecosystem—the interactive Early ecology was driven by a it relies on geology, geomorphology,
relationship between living desire for knowledge. Later, it was climatology, chemistry, physics,
organisms and the environment used to find better ways to exploit genetics, sociology, and more.
in which they live. Later ecologists the natural world for human needs. Ecology influences local and
developed mathematical models to As time went on, the consequences national government decisions
forecast population dynamics within of this exploitation became about urbanization, transportation,
ecosystems. Evolutionary theories increasingly evident. Deforestation industry, and economic growth.
also advanced with the discovery was highlighted as a problem as The challenges posed by climate
of the structure of DNA, and the early as the 18th century, and the change, rising sea levels, habitat
evolutionary “vehicle” provided problems of air and water pollution destruction, the extinction of
by mutation as DNA is replicated. became obvious in industrialized species, plastic and other forms of
nations in the 19th century. In 1962, pollution, and a looming water crisis
New frontiers Rachel Carson’s book Silent Spring pose serious threats to human
Improved technology opened up alerted the world to the dangers of civilization. They demand radical
new possibilities for ecology. An pesticides, and six years later Gene policy responses based on sound
electron microscope can now Likens demonstrated the link science. Ecology will provide the
make images to half the width of a between power station emissions, answers. It is up to governments
hydrogen atom, and computer acid rain, and fish deaths. to apply them. ■
programs can analyze the sounds In 1985, a team of Antarctic
made by bats and whales, which are scientists discovered the dramatic
higher or lower than can be heard depletion of atmospheric ozone
by the human ear. Camera traps and over Antarctica. The link between
infrared detectors photograph and greenhouse gases and a warming
film nocturnal creatures, and tiny of Earth’s lower atmosphere had
satellite devices fitted to birds can been made as early as 1947 by Even in the vast and
track their movements. G. Evelyn Hutchinson, but it was mysterious reaches of the sea
In the laboratory, analysis of decades before there was a scientific we are brought back to the
the DNA of feces, fur, or feathers consensus on the man-made causes fundamental truth that
indicates which species an animal of climate change. nothing lives to itself.
belongs to, and throws light on Rachel Carson
the relationship between different The future
organisms. It is now easier than Modern ecology has come a long
ever for ecologists to collect data, way since the science was first
helped by a growing army of recognized. It now draws on many
citizen scientists. disciplines. In addition to zoology,
THE STO
OF EVOL
RY
UTION
18 INTRODUCTION
HMS Beagle sets sail on a

James Hutton presents In his Essay on the Theory circumnavigation of the world, with
his theory that Earth is of the Earth, Georges Cuvier Charles Darwin serving as the
much older than was suggests that fossils are the voyage’s naturalist. The trip provides
previously believed, and remains of extinct creatures Darwin with the information that
that Earth’s crust is wiped out by periodic inspired his theory of evolution
continuously changing. “catastrophic” events. by natural selection.
1785 1813 1831
1809 1823
Jean-Baptiste Lamarck Amateur fossil

publishes Philosophie Zoologique, hunter Mary Anning
where he argues that animals acquire uncovers the first intact
characteristics as a consequence of plesiosaurus skeleton.
use or nonuse of different body parts,
triggering mutations over generations.
A
ncient myths, religions, and was the driving force behind this these processes take place slowly,
philosophies all reflect an change. He speculated that Earth’s history had to be much
enduring fascination with characteristics acquired by animals longer than was previously thought.
how the world began and man’s during their lifetime were inherited
place in the story of life on Earth. In by the next generation: giraffes, for Natural selection
the West, Christianity held that all example, became slightly longer- In 1858, Charles Darwin and Alfred
animals and plants were the result necked by stretching up to reach Russel Wallace delivered a paper
of a perfect creation. On the chain higher leaves, and passed this trait that would change biology forever.
or ladder of being, no species could to their offspring; over many Darwin’s observations on the epic
ever move from one position to generations, giraffes grew longer voyage of the Beagle (1831–36),
another. Species were immutable, and longer necks. his correspondence with other
an idea called essentialism. Fossil evidence of extinct life naturalists, and the influence
The 18th-century Age of forms with features that resembled of Thomas Malthus’s writings
Enlightenment began to challenge modern descendants, found by inspired Darwin’s insight that
orthodox Christian beliefs. French pioneering geologists such as evolution came about by what he
zoologist Jean-Baptiste Lamarck Georges Cuvier, also suggested called natural selection. He spent
rejected the prevailing Bible-based Earth had more ancient origins. 20 years gathering supporting data,
notion of Earth being only a few Meanwhile James Hutton and but when Wallace wrote to him
thousand years old. He argued that Charles Lyell argued that geological with the same idea, Darwin
organisms must have changed from features could be accounted for by realized it was time to go public.
simple life forms to more complex the constant, ongoing processes His subsequent book, On the
ones over millions of years, and that of erosion, and deposition—a view Origin of Species by Means of
the “transmutation” of species called uniformitarianism. Because Natural Selection, provoked outrage.
THE STORY OF EVOLUTION 19
Gregor Mendel’s paper The Selfish Gene by

“Experiments with Plant Hybrids” evolutionary biologist Richard
outlines findings from his pea Dawkins offers a new
plant experiments, laying perspective on evolution,
the foundations for the looking at the gene, as opposed
field of genetics. to the species or group.
1866 1976
1859 1953 2003
Darwin elaborates on his In The Eagle pub in The Human Genome

theories of evolution in On the Cambridge, UK, Crick and Project produces the first
Origin of Species by Means Watson announce that genetic blueprint of
of Natural Selection, which they have discovered Homo sapiens.
is an instant sellout. the structure of DNA.
Although the idea of evolution of thought were complementary, that genetic information is “written”
became widely accepted, the rather than contradictory. In 1942, on DNA molecules. The errors that
mechanism that made natural Julian Huxley articulated the occur when DNA copies itself create
selection possible was not yet synthesis between Mendel’s mutations—the raw materials for
known. In 1866, an Austrian monk genetics and Darwin’s theory evolution. By the 1980s it was
called Gregor Mendel made a huge of natural selection in his book possible to map and manipulate the
contribution to genetics when he Evolution: The Modern Synthesis. genes of individuals and species. In
published his findings on heredity the 1990s, the mapping the human
in pea plants. Mendel described The double helix genome paved the way for medical
how dominant and recessive traits Advances in technology such as research into gene therapy.
pass from one generation to the X-ray crystallography led to more Ecologists also want to establish
next, by means of invisible “factors” discoveries in the 1940s and ’50s, whether genes influence behavior.
that we now call genes. and the foundation of the new Back in 1964, William D. Hamilton
The rediscovery of Mendel’s discipline of molecular biology. popularized the concept of genetic
work in 1900 initially sparked sharp In 1944, chemist Oswald Avery relatedness (“kin selection”) to
debate between his supporters and identified deoxyribonucleic acid explain altruistic behavior in
many Darwinians. At the time, (DNA) as the agent for heredity. animals. In The Selfish Gene (1976),
evolution was believed to be based Rosalind Franklin and Raymond Richard Dawkins further advanced
on the selection of small, blending Gosling photographed strands of the the gene-centered approach. It is
variations, but Mendel’s variations DNA molecule in 1952, and James clear that aspects of evolutionary
clearly did not blend. Three decades Watson and Francis Crick confirmed biology will still spark debate as
later, geneticist Ronald Fisher and its double helix structure the long as ecologists continue to
others argued that the two schools following year. Crick then showed develop Darwin’s theory. ■
20
TIME IS INSIGNIFICANT,
AND NEVER A DIFFICULTY
FOR NATURE
EARLY THEORIES OF EVOLUTION
B
efore the 18th century, most material, struck off the Sun by a
IN CONTEXT people believed that plant comet, that had taken 70,000 years
and animal species stayed to cool (a huge underestimate, in
KEY FIGURES
unchanged throughout time—a view fact). As Earth cooled, species had
The Comte de Buffon
now known as essentialism. This appeared, died off, and were finally
(1707–88), Jean-Baptiste
idea came under challenge as a replaced by ancestors of those
Lamarck (1744–1829) result of two developments: the known today. Noting similarities
BEFORE intellectual movement known as the among animals such as lions,
1735 Swedish botanist Carl Enlightenment (c. 1715–1800), and tigers, and cats, Buffon deduced
Linnaeus publishes Systema the Industrial Revolution (1760–1840). that 200 species of quadrupeds had
Naturae, a system of biological The Enlightenment was marked evolved from just 38 ancestors. He
classification that later helped by scientific progress and increased also believed that changes in body
to determine species’ ancestry. questioning of religious orthodoxy, shape and size in related species
such as the claim that God created had occurred in response to living
1751 In “Système de la nature” Earth and all living things in seven in different environments.
French philosopher Pierre days. Then, as the Industrial In 1800, French naturalist Jean-
Louis Moreau de Maupertuis Revolution gathered pace, canals, Baptiste Lamarck went further. In a
introduces the idea that railroads, mines, and quarries lecture at the Museum of Natural
features can be inherited. cut through rock strata and revealed
thousands of fossils, mostly of
AFTER animal and plant species that no
1831 Etienne Geoffroy Saint- longer existed and had never been
Hilaire writes that sudden seen before. These suggested that
environmental change can life began long before the widely
cause a new species to develop accepted creation date of 4400 bce, Nature is the system of laws
from an existing organism. deduced from biblical sources. established by the Creator for
the existence of things and
1844 In Vestiges of the Natural Animal adaptation for the succession of creatures.
History of Creation, Scottish In the late 1700s, French scientist The Comte de Buffon
geologist Robert Chambers Georges-Louis Leclerc, Comte de
argues—anonymously—that Buffon, upset church authorities
simple creatures have evolved by asserting that Earth was much
into more complex species. older than the Bible suggested. He
believed it was formed from molten

See also: Extinction and change 22 ■ Uniformitarianism 23 ■ Evolution by
natural selection 24–31 ■ The rules of heredity 32–33
History in Paris, he argued that

traits acquired by a creature during
its lifetime could be inherited by
its offspring—and that a buildup
of such changes over many
generations could radically alter …continuous use of
an animal’s anatomy. any organ gradually
Lamarck wrote several books strengthens, develops
in which he developed this idea and enlarges that organ.
of transmutation. He argued, for Jean-Baptiste Lamarck
instance, that the use or nonuse of Jean-Baptiste
body parts eventually resulted in
Lamarck
such features becoming stronger,
weaker, bigger, or smaller in a Born in 1744, Jean-Baptiste
species. For example, the ancestors Lamarck attended a Jesuit
of moles probably had good college before joining the
eyesight, but over generations develop from simple to more French army. Forced by an
this deteriorated because moles did complex forms in a “ladder” of injury to resign, he studied
not require vision as they burrowed progress. The other, via the medicine and then pursued
underground. Similarly, giraffes inheritance of acquired traits, his passion for plants, working
gradually developed longer necks helped them adapt better to their at the Jardin du Roi (Royal
to enable them to reach leaves environment. When Charles Darwin Garden) in Paris. Supported
growing high up in trees. developed his theory of evolution by the Comte de Buffon,
by means of natural selection, he Lamarck was elected to the
Academy of Sciences in 1779.
Drivers of evolution would reject many of Lamarck’s
When the Jardin’s main
Larmarck’s ideas about inherited ideas, but both men shared the building became the new
acquired traits were part of a wider belief that complex life evolved National Museum of Natural
early theory of evolution. He also over an immense period of time. ■ History during the French
believed that the earliest, simplest Revolution (1789–99), Lamarck
forms of life had emerged directly was placed in charge of the
Fossil finds changed ideas about
from nonliving matter. Lamarck how life began. The first example of an study of insects, worms, and
identified two main “life forces” articulated plesiosaur—Plesiosaurus microscopic organisms. He
driving evolutionary change. One, dolichodeirus—was discovered in 1823 coined the biological term
he believed, made organisms by Mary Anning in Dorset, England. “invertebrate” and often used
the relatively simpler forms of
such species to illustrate his
“ladder” of evolutionary
progress. However, Lamarck’s
work was controversial and
he died in poverty in 1829.
Key works
1802 Research on the

Organization of Living Bodies
1809 Zoological Philosophy
1815–22 Natural History of
Invertebrate Animals
22
A WORLD PREVIOUS
TO OURS, DESTROYED
BY CATASTROPHE
EXTINCTION AND CHANGE
I
n the early days of studying believe that the evidence of fossil
IN CONTEXT fossils, many people denied remains supported a theory of
they could be extinct species. evolution. Nevertheless, Cuvier’s
KEY FIGURE
They failed to see why God would central views have continued to
Georges Cuvier (1769–1832)
create and destroy creatures before win support, and modern evidence
BEFORE humans ever appeared, arguing points to at least five catastrophic
Late 1400s Leonardo da Vinci that unfamiliar fossil species might mass extinction events in Earth’s
argues that fossils are the still be living somewhere on Earth. past, including the one that wiped
remains of living creatures, In the late 18th century, French out the dinosaurs. Unlike Cuvier,
not just shapes spontaneously zoologist Georges Cuvier looked however, today’s scientists know
formed in the earth. into this by exploring the anatomy that life is not recreated out of
of living and fossil elephants. He nothing after a catastrophe. Rather,
1660s English scientist Robert proved that fossil forms such as when a mass extinction event kills
Hooke suggests that fossils are mammoths and mastodons were off many species, those left will
extinct creatures, since no anatomically distinct from living evolve and multiply—sometimes
similar forms can be found elephants, so they must represent relatively quickly—to fill vacant
on Earth today. extinct species. (It was highly ecological niches, as the mammals
unlikely that they still lived on did after the age of the dinosaurs. ■
AFTER Earth without being noticed.)
1841 English anatomist Cuvier believed that Earth had
Richard Owen calls huge experienced a series of distinct
reptile fossils “dinosaurs.” ages, each of which ended with a
“revolution” that destroyed existing
1859 Charles Darwin’s On the
flora and fauna. He did not, though,
Origin of Species explains how
evolution can occur through
“natural selection.” Cuvier coined the name “mastodon”
for its Greek meaning of “breast tooth,”
1980 US scientists Luis referring to the nipplelike patterns on
and Walter Alvarez present the creature’s teeth, which were unlike
evidence that an asteroid those of any living elephants.
hit Earth at the time of the
extinction of the dinosaurs. See also: Evolution by natural selection 24–31 ■ Ecological niches 50–51
■ An ancient ice age 198–199 ■ Mass extinctions 218–223
NO VESTIGE OF A
BEGINNING, NO
PROSPECT
UNIFORMITARIANISM
OF AN END
U
niformitarianism is the
IN CONTEXT theory that geological
processes, such as the
KEY FIGURE
laying down of sediment, erosion,
James Hutton (1726–97)
and volcanic activity, occur at the
BEFORE same rate now as they did in the … from what has actually
1778 The Comte de Buffon, a past. The idea emerged in the late been, we have data for
French naturalist, suggests 18th century, as mining, quarrying, concluding [what] is
that Earth is at least 75,000 and increased travel brought ever to happen thereafter.
years old—far older than most more geological features to light, James Hutton
people believed at the time. including unusual rock strata and
previously unknown fossils, whose
1787 German geologist origins were then widely debated.
Abraham Werner proposes The generally accepted view
that Earth’s layers of rock that Earth was only a few thousand
formed from a great ocean that years old had been challenged by
once covered the entire planet. the Comte de Buffon, and in 1785 that most geological processes
His followers became known Scottish geologist James Hutton happen so gradually that the
as Neptunists. also argued for Earth’s far greater features he was discovering must
antiquity. Hutton’s ideas were be astronomically old.
AFTER formed during expeditions around Uniformitarianism was not
1802 James Hutton’s theory Scotland to examine layers of rock. generally accepted at once, not
of uniformitarianism reaches a He believed that Earth’s crust was least because it challenged a literal
wider audience when Scottish constantly changing, albeit mostly interpretation of the creation stories
geologist John Playfair slowly, and could see no reason to of the Old Testament. However, a
publishes Illustrations of the suggest that the complex geological new generation of geologists, such
Huttonian Theory of the Earth. actions of layering, erosion, and as John Playfair and Charles Lyell,
uplifting took place faster in the threw their intellectual weight
1830–33 Principles of Geology, distant past than they did in the behind Hutton’s ideas, which also
by Scottish geologist Charles present. Hutton also understood inspired a young Charles Darwin. ■
Lyell, supports and builds on
the uniformitarian ideas of See also: Early theories of evolution 20–21 ■ Evolution by natural selection 24–31
James Hutton. ■ Moving continents and evolution 212–213 ■ Mass extinctions 218–223
THE STRUGGLE FOR

EXISTENCE
EVOLUTION BY NATURAL SELECTION
26 EVOLUTION BY NATURAL SELECTION
N
atural selection, a concept
IN CONTEXT developed by British
naturalist Charles Darwin
KEY FIGURE
and set out in his book On the
Charles Darwin (1809–82)
Origin of Species by Means of
BEFORE Natural Selection (1859), is the Natural selection is daily
1788 In France, Georges-Louis key mechanism of evolution in and hourly scrutinizing,
Leclerc, Comte de Buffon, organisms, resulting in different throughout the world,
completes his 36-volume survival rates and reproductive the slightest variations.
Histoire Naturelle, outlining abilities. Those organisms that have Charles Darwin
early ideas about evolution. higher breeding success pass on
their genes to more of the next
1809 Jean-Baptiste Lamarck generation, so individuals with
proposes that creatures evolve these characteristics become
by inheriting acquired traits. more common.
AFTER To the Galapagos thousands of years. As Darwin
1869 Friedrich Miescher, a The young Charles Darwin first looked at landscapes around the
Swiss doctor, discovers DNA, began to consider evolution during world that had been affected by
although its genetic role is not his pioneering scientific expedition processes of erosion, deposition, and
yet understood. around the world aboard HMS volcanism, he began to speculate
1900 The laws of inheritance Beagle from 1831 to 1836. As a young about animal species changing over
man, Darwin accepted the orthodox very long time periods, and the
based on the pea plant
interpretation of the Bible, that Earth reasons for such changes. By
experiments of Austrian
was only a few thousand years old. examining fossils and observing
scientist Gregor Mendel in the However, while he was on board living animals, Darwin identified
mid-1800s are rediscovered. the Beagle, Darwin read Scottish patterns; he noticed, for example,
1942 British biologist Julian geologist Charles Lyell’s recently that extinct species had often been
Huxley coins the term “modern published Principles of Geology, in replaced by similar, but distinct,
synthesis” for the mechanisms which Lyell demonstrated that rocks modern ones.
thought to produce evolution. bore traces of tiny, gradual, and Darwin’s field work on the
cumulative change over vast time islands of the Galapagos archipelago
periods—millions, rather than off South America in the fall
Charles Darwin Born in Shropshire, UK, in 1809, instantly. Despite continuing

Darwin was fascinated by natural ill-health, Darwin fathered
history from a young age. While 10 children and never stopped
at Cambridge University, he studying and developing new
became friendly with several theories. He died in 1882.
influential naturalists, including
John Stevens Henslow. As a result,
Darwin was invited to join the Key works
HMS Beagle expedition around the
world. Henslow helped Darwin 1839 Zoology of the Voyage
catalog and publicize his finds. of HMS Beagle
Darwin’s research brought him 1859 On the Origin of Species
fame and recognition—the Royal by Means of Natural Selection
Society’s Royal Medal in 1853, 1868 The Variation of Animals
nd fellowship of the Linnean and Plants under Domestication
Society in 1854. In 1859, his book 1872 The Expression of
On the Origin of Species sold out Emotions in Man and Animals

See also: Early theories of evolution 20–21 ■ The rules of heredity 32–33 ■ The role of DNA 34–37 ■ The selfish gene 38–39
■ The food chain 132–133 ■ Mass extinctions 218–223 ■ Population viability analysis 312–315
of 1835 provided especially strong Comparison of Galapagos finch

evidence for his later theory of bill structure
evolution by natural selection. Here,
he observed that the shape of the
carapaces (shells) of giant tortoises
varied slightly from island to island.
Darwin was also intrigued to find
that there were four broadly similar,
yet clearly distinct, varieties of
mockingbirds, but that no single
island had more than one species Geospiza magnirostris Geospiza fortis
The short, sharp bill of the Large The bill of the Medium Ground Finch
of the bird. He saw small birds, Ground Finch, the biggest of Darwin’s is variable, evolving rapidly to adapt
too, that looked alike but had a finches, enables it to crack nuts. to whatever size seeds are available.
range of beak sizes and shapes.
Darwin deduced that each group
possessed a common ancestor but
had developed diverse traits in
different environments.
Darwin’s conclusions
On Darwin’s return to England, the Geospiza parvula Certhidea olivacea
differing beaks of the small birds The stubby bill of the Small Tree Finch, The slender, probing bill of the Green
which forages in foliage, suits its diet Warbler-finch helps it catch small
he had found on the Galapagos, of seeds, fruits, and insects. insects and spiders.
usually called “finches” although
they are not in the true finch family,
set him thinking. He knew that populations had evolved in different Malthus predicted that population
a bird’s beak is its key tool for Galapagos habitats, each group growth would eventually outstrip
feeding, so its length and shape adapted for a more or less specialist food production. This idea matched
offer clues to its diet. Later research diet by a process that he would the evidence Darwin had observed
revealed that there are 14 different later call “natural selection.” Over of ongoing competition between
finch species on the Galapagos time, the finch populations had individual animals and species for
islands. The differences in their become distinct species. resources. This competitive aspect
beaks are marked and significant. In the early 21st century, formed the backbone of Darwin’s
For example, cactus finches have researchers at Harvard University coalescing theory of evolution.
long, pointed beaks that are ideal uncovered new evidence of how By 1839, Darwin had developed
for picking seeds out of cactus this happens at a genetic level. an idea of evolution by natural
fruits, while ground finches have Their findings, published in 2006, selection. He was, though, reluctant
shorter, stouter beaks that are showed that a molecule called to publish because he understood
better suited for eating large seeds calmodulin regulates the genes that the theory would unleash a
on the ground. Warbler finches have involved in shaping birds’ beaks, storm of controversy from those
slender, sharp beaks, which are and is found at higher levels in who would view it as an attack
ideal for catching flying insects. longer-beaked cactus finches than on religion and the Church. When,
Darwin speculated that the in shorter-beaked ground finches. in 1857, he began receiving
finches were descended from a communications from fellow British
common ancestral finch that had Refining the theory naturalist Alfred Russel Wallace,
reached the archipelago from the Darwin was influenced by Thomas who had independently arrived at
mainland of South America. He Malthus’s An Essay on the Principle very similar conclusions, Darwin
concluded that a variety of finch of Population (1798), in which realized he had to publish his ❯❯

ideas. Papers by Darwin and Sex (1871). This theory was distinct
Wallace were jointly presented from natural selection, as Darwin
at a meeting of the Linnean recognized that animals select
Society of London in July 1858, mates based on characteristics that
under the title “On the Tendency do not simply favor survival. For
of Species to form Varieties; and I see no good reasons why example, when Darwin considered
on the Perpetuation of Varieties the views given in this the spectacular but cumbersome
and Species by Natural Means volume should shock the tails of male peafowl (peacocks), he
of Selection.. religious views of anyone. could not imagine the tail playing
The following year, Darwin Charles Darwin any role in helping the individual
published the theory in On the bird to survive. He concluded that
Origin of Species by Means of they were designed to boost an
Natural Selection. It offended some individual’s chance of reproductive
scientists because it differed from success. Peahens choose males
Lamarck’s ideas of transmutation, with the brightest tails, so the
and also upset creationists who genetic material of these showy
argued that it undermined a literal helped an individual organism males is passed to the next
interpretation of the Bible. Others live longer and reproduce more generation. Bright tail feathers
felt that the theory did not account successfully would be passed on indicate that the bird is healthy, so
for the huge range of characteristics to more offspring, while those that choosing a mate with a bright tail
in species and called it “unguided” made the organism less successful is a good strategy for the peahen.
and “nonprogressive.” would be lost. Darwin called this However, Darwin’s idea that
Darwin was confident. He knew “natural selection”—a process females choose a mate came under
that all individual organisms in a that, over generations, enabled fire; 19th-century society could
species show a degree of natural a population of any given species accept that males competed to
variation; some have longer to adapt better and thrive in its reproduce (intrasexual selection),
whiskers, or shorter legs, or brighter chosen habitat. but intersexual selection, where
colors, for instance. Because one sex (usually the female) makes
members of all species compete for Sexual selection the choice, was ridiculed.
limited resources, he deduced that Darwin also developed a theory Reproductive success is clearly
those whose traits are best suited of sexual selection. First outlined in essential for the future of a species.
to their environment are more likely On the Origin of Species, this was Natural selection is often described
to survive and reproduce. He also developed further in The Descent as “survival of the fittest,” but
argued that characteristics that of Man, and Selection in Relation to longevity alone is not particularly
Natural selection
There is variation There is differential reproduction. There is heredity. End result:

in traits. No environment can support unlimited The dark beetles have If darkness is the
For example, some population growth, so some individuals more dark offspring winning trait, producing
beetles are pale and lose out. Here, birds eat the pale because this trait has a more offspring, in time,
others dark. beetles, so fewer of them reproduce. genetic basis. all beetles will be dark.

Kin selection
The term “kin selection” was
first used by British biologist
John Maynard Smith in 1964.
It is the evolutionary strategy
that favors the reproductive
success of an organism’s
relatives, prioritizing them
above the individual’s own
survival and reproduction.
It occurs when an organism
engages in self-sacrificial
behavior that benefits its
relatives. Charles Darwin was
the first to discuss the concept
when he wrote about the
apparent paradox represented
by altruistic nonbreeding
social insects, such as worker
honeybees, which leave
reproduction to their mothers.
British evolutionary biologist
William Donald Hamilton
proposed that bees, for
example, behave in an
altruistic manner—assisting
The peacock with the most splendid description of the scientific process. others in reproduction—when
tail will attract the most peahens. Its In 1930, British geneticist Ronald the genetic closeness of the
bright tail will be passed on to its male Fisher wrote The Genetical Theory two bees and the benefit to
offspring, which will find it similarly the recipient outweigh the
easy to attract mates.
of Natural Selection, which
combined Darwin’s theory of cost of altruism to the giver.
natural selection with the ideas This is called Hamilton’s Rule.
helpful. If individual A lives 10 of heredity that the 19th-century
times as long as individual B, but Austrian scientist Gregor Mendel
the latter produces twice as many had developed. In 1937, Ukrainian–
offspring that then also breed, B American geneticist Theodosius ❯❯
will pass on more genes to the next
generation than the longer-lived A.
Building on the theory

Many of Darwin’s and Wallace’s
ideas have proved remarkably
accurate, despite the fact that Why do some die and some
the workings of genetics were not live?… the answer was
understood at the time. Although clearly, that on the whole the
Darwin himself had used the best fitted live.
term “genetic” as an adjective Alfred Russel Wallace
to describe the as-yet-unknown In honeybee colonies, female
mechanism of inheritance, it was worker bees look after the queen
bee. They build the honeycomb,
British biologist William Bateson, gather nectar and pollen, and feed
in the early 20th century, who first larvae, but they do not breed.
used the term “genetics” in a
Albinism, as in this albino leopard organism carrying it, whereas environment, they become more
gecko, is a mutation causing a lack another might affect all its offspring common over the course of
of pigment. This mutation hinders the and future generations. generations. Over time, they may
gecko’s chances of survival, making it
lighter colored and sensitive to light.
Inherited mutations may or may produce large enough divergences
not alter an individual’s phenotype – from the parent population for a
its physical traits and behavior. If new species to evolve—a process
Dobzhansky put forward the idea mutations do affect the phenotype, called speciation.
that regularly occurring genetic they may be to its advantage or Mutation rates are usually very
mutations are sufficient to provide disadvantage, helping or hindering low, but the process is ever-present.
the genetic diversity—and an organism’s ability to survive The changes may be beneficial,
therefore different traits—that and reproduce successfully. If they neutral, or harmful. They do not
makes natural selection possible. hinder, they are likely to disappear occur in response to an organism’s
He wrote that evolution was a from the population; if they help needs, and are, in that respect,
change in the frequency of an an organism adapt better to its random. However, some types of
“allele” in the gene pool, an allele mutations occur more frequently
being one of the alternative forms of than others. Scientists now know,
a gene that arise by mutation. for example, that evolution can take
A mutation is a permanent place very rapidly in bacteria
alteration in the sequence of because of their frequent mutations.
deoxyribonucleic acid (DNA), the
molecule that makes up a gene The vast majority of large Different rates of evolution
in one individual, resulting in mutations are deleterious; The ancestors of all life on Earth
a sequence that differs from that small mutations are both far were very simple organisms.
of other members of the species. more frequent and more likely Recent scientific research suggest
Mutations may occur as the result to be useful. that the earliest “biogenic” rocks—
of the miscopying of DNA during Ronald Fisher derived from early life forms—date
cell division, or they may be caused back nearly four billion years. In
by environmental factors, such as that time, highly complex life forms
damage resulting from the sun’s have evolved, and later fossils
ultraviolet radiation. One mutation of species that look more similar
might affect only the individual to those of today reveal what has

Evolution in real time
Richard Lenski, a professor
Individuals within at Michigan State University,
a species have established the Long-term
a variety of forms Experimental Evolution
Seen in the light of evolution, of a characteristic. project in 1988. For more than
biology is, perhaps, 25 years, he studied 59,000
intellectually the generations of the E. coli
most satisfying and bacterium. During this time,
inspiring science. he observed that the species
Theodosius Dobzhansky used the glucose solution
it lived in more efficiently,
increasing in size but also
The individuals with
growing faster. Also, a new
the characteristic best
species had evolved that was
suited to the environment
able to use a compound in the
are more likely to survive
solution called citrate, which
and breed.
the parent bacterium could
occurred. For example, a fossil
not. Evolving bacteria can
record stretches back 60 million
pose a potential threat to
years for ancestors of the horse. humans. Increasing antibiotic
The earliest of these were dog- use destroys many disease-
sized forest-dwelling animals with causing bacteria, but not those
several toes on each foot. Evolution with mutations that make
produced much larger horses with These them resistant to the drugs.
just a single hoof on each foot, characteristics As the non-resistant bacteria
adapted for life on open grasslands are passed on are killed off, the resistant
where they would often have had to the next strains become more
to outrun predators. generation. dominant, multiplying and
Peppered moths (biston passing on their mutations
betularia) reveal change over a to future generations. That
shorter period. The moth is usually is natural selection at work.
pale, providing camouflage against
the bark of birch trees, but a
mutation produces some black
moths. Before the 19th century,
most peppered moths were pale.
During the Industrial Revolution
(1760–1840), however, smoky air left
deposits of soot on trees and
buildings in British cities, and the
black form became much commoner.
By 1895, 95 percent of peppered
moths in Britain’s cities were black,
as paler moths were eaten by birds
because their coloring provided no
camouflage. This phenomenon
continues to act as an example of Escherichia (E.) coli bacteria
Two peppered moths exhibit can cause serious gut and other
Darwin’s theory in action today, as evolution at work, the lower one an infections that will be increasingly
the pale moth becomes common example of industrial melanism. The difficult to treat as drug-resistant
once more due to the declining soot dark variety began to appear in British strains of E. coli multiply.
concentrations in Britain’s cities. ■ cities in the early 1800s.
32
HUMAN BEINGS
ARE ULTIMATELY
NOTHING BUT
CARRIERS FOR
THE RULES OF HEREDITY
GENES
L
ong before scientists was not the case when he was
IN CONTEXT cracked the genetic code, working in his monastery garden.
in 1866 an Austrian monk When he crossed a plant that
KEY ECOLOGIST
named Gregor Mendel was the first always produced green peas with
Gregor Mendel (1822–84)
to show how traits are transferred one that always produced yellow
BEFORE through the generations. By means peas, the result was not yellowish-
1802 French biologist Jean- of much painstaking research, green peas—instead, all the peas
Baptiste Lamarck suggests Mendel accurately predicted the were yellow.
that traits acquired during the basic laws of inheritance.
lifetime of an organism are When Mendel began his Mendel’s labors
transmitted to its offspring. experiments, scientists believed During the course of his research
that the various traits seen in (1856–63), Mendel grew nearly
1859 Charles Darwin proposes plants and animals were handed 30,000 pea plants over several
his theory of evolution and down through a “blending” process. generations and carefully recorded
natural selection in his book However, Mendel noticed that this the results. He focused on traits
On the Origin of Species by
Means of Natural Selection.
Mendel’s pea experiment
AFTER
Mendel’s
1869 Swiss chemist Friedrich experiment
Miescher identifies DNA, with growing peas PARENT GENERATION
which he terms “nuclein.” proved that the gene
carrying the yellow 1 green 1 yellow
1953 Molecular biologists— coloration was
including Briton Francis dominant while the
Crick and American James gene for green was all yellow
Watson—discover the recessive.
F1 GENERATION
structure of DNA.
2000s Researchers in the
field of epigenetics describe
inheritance by mechanisms
other than through the DNA F2 GENERATION
sequence of genes.
1 green 3 yellows

See also: Early theories of evolution 20–21 ■ Evolution by natural selection 24–31
■ The role of DNA 34–37 ■ The selfish gene 38–39
dominant or recessive. When both

inherited factors are dominant,
the resulting plant will show the
dominant trait. With a pair of
recessive factors, the plant will
Heredity provides show the recessive trait. However,
for the modification if one dominant and one recessive
of its own machinery. factor are present, the plant will
James Mark Baldwin show the dominant trait.
American psychologist
Pioneering geneticist Gregor Johann Mendel

Mendel published his paper in
1866, but no one took much notice Born Johann Mendel in 1822
until 1900, when the botanists on a farm in Silesia—then part
Hugo de Vries, Carl Erich Correns, of the Austrian Empire and
(phenotypes) that had only two and Erich Tschermak von now in the Czech Republic—
distinct forms—for example, white Seysenegg discovered his work. Mendel studied philosophy
or purple flowers. When examining Scientists then began proving and physics at the University
the trait of yellow or green peas, Mendel’s theories more widely. of Olomouc (1840–43). At this
Mendel took green pea plants and Within just ten years, scientists time, he became interested
cross-pollinated them with yellow named the pairs of factors “genes” in the work of Johann Karl
pea plants. The peas produced from and showed that they are linked on Nestler, who was researching
hereditary traits in plants
this parent generation were all yellow chromosomes. It is now known that
and animals. In 1847 Mendel
and Mendel named them the F1 inheritance is far more complex
entered a monastery, where
generation. He then cross-pollinated than Mendel recognized, but his he was given the name
pea plants from the F1 generation meticulous research continues to Gregor. He then went on
with each other to produce the F2 form the basis for modern studies. ■ to study science further at
generation. He found that some Vienna University (1851–53).
peas produced were yellow and When Mendel returned
some were green. The F1 generation to his monastery in 1853,
showed only one trait (yellow), which the abbot Cyril Napp gave
Mendel called “dominant.” However, him permission to use the
in the F2 generation 75 percent had gardens for his research into
the dominant yellow trait and 25 hybridization. Mendel himself
percent displayed the nondominant became an abbot in 1868 and
—or “recessive”—green trait. no longer had time for his
experiments. Although he
never received credit for his
Laws of inheritance discoveries during his lifetime,
Mendel theorized that every pea he is widely regarded as the
plant has two factors controlling founder of modern genetics.
each trait. When plants are cross-
pollinated, one factor is inherited Key works
from each plant. A factor can be
1866 “Experiments with Plant
Hybrids,” Verhandlungen des
Pea plants provided the raw data naturforschenden Vereines
that Mendel used to develop his in Brünn
theories explaining the transmission of
traits from one generation to the next.
34
IN CONTEXT
WE’VE
KEY FIGURES
Francis Crick (1916–2004),
Rosalind Franklin (1920–
58), James Watson (1928–),
DISCOVERED
Maurice Wilkins (1916–2004)
BEFORE
1910–29 US biochemist
Phoebus Levene describes the
THE SECRET
chemical components of DNA.
1944 US researchers Oswald
Avery, Colin Macleod, and
Maclyn McCarty show that
OF LIFE
DNA determines inheritance.
AFTER
1990 British researchers,
led by embryologist Ian
Wilmut, successfully clone
THE ROLE OF DNA an adult mammal—a sheep
named Dolly.
2003 Scientists complete
the mapping of the entire
human genome.
T
he discovery of the structure
of DNA (deoxyribonucleic
acid) in 1953 is one of
the most important scientific
breakthroughs to date. It offered
the key to understanding the very
building blocks of life and explained
how genetic information is stored
and transferred. Englishman Francis
Crick and American James Watson
famously celebrated their joint
discovery in a low-key fashion
at their local pub in Cambridge,
followed by a letter published in
the journal Nature. Their discovery
had enormous potential for scientific
advances and had an important
impact on many fields of research,
from medicine to forensic science,
taxonomy, and agriculture. The
ramifications of their work still
reverberate today, as methods of

See also: Early theories of evolution 20–21 ■ Evolution by natural selection 24–31 ■ The rules of heredity 32–33 ■ The
selfish gene 38–39 ■ A system for identifying all nature’s organisms 86–87 ■ Biological species concept 88–89
College, London, Franklin and

Wilkins were developing methods of
X-raying DNA to view its structure.
Watson had seen examples of
Franklin’s work that hinted at DNA’s
helical shape shortly before he and DNA is like a computer
Crick announced their breakthrough. program but far, far more
In 1962 Crick, Watson, and advanced than any
Wilkins were awarded the Nobel software ever created.
Prize for Physiology or Medicine. Bill Gates
Franklin, who died in 1958, never
received recognition for her part
in the discovery during her lifetime,
Molecular biologists James Watson although Crick and Watson openly
(left) and Francis Crick (right), pictured acknowledged that her work was
in 1953 with their double helix model essential to their success.
of DNA. Watson called DNA “the most with T to form base pair AT, and
interesting molecule in all nature.”
Double helix structure G always pairs with C to form
DNA is a molecule featuring two base pair GC.
handling genetic material advance long, thin strands that coil around DNA is the blueprint for life.
and we learn more about how each other to resemble a twisted Sequences of bases along the
individual genes operate. ladder, in a shape known as a DNA strand constitute the genes
Crick and Watson’s breakthrough double helix. Using the ladder that provide the information that
was the culmination of decades analogy, the sides of the ladder are determines the complete form and
of research by numerous scientists, made up of deoxyribose (a sugar) physiology of an organism. A triplet
including Rosalind Franklin and and phosphate, while the rungs of bases is known as a codon, and
Maurice Wilkins. While Crick and of the ladder consist of paired each codon specifies the production
Watson worked with 3-D models nitrogenous bases, adenine (A), of one of 20 amino acids; the order
to figure out how the components guanine (G), cytosine (C), and in which the amino acids join
of DNA fitted together, at King’s thymine (T). A always pairs up together in a chain determines ❯❯
Genetic engineering useful—have greatly simplified

and accelerated the process. In
Understanding the structure theory, geneticists can now splice
of DNA has enabled scientists any gene with any other. They
to change or “engineer” the have attempted some intriguing
genetic material in cells. It is combinations, such as the
possible to cut out a gene from insertion of the gene for producing
one organism (the donor) and spider silk into goat DNA so
place it into the DNA of another that goats produce milk rich in
organism. When this practice proteins. Other substances that
was first attempted in the 1970s can be produced by modifying
it was both difficult and time- genes are hormones and vaccines.
consuming, but technological In gene therapy, a genetically
advances—such as Clustered modified vector (often a virus) is A scientist analyzes a sample
Regularly Interspaced Short used to carry a gene into the DNA of DNA. Genetic manipulation in
Palindromic Repeats, or CRISPR, of an organism to replace a faulty medicine is standard practice and
which has been particularly or unwanted gene. DNA profiling is a vital forensic tool.
36 THE ROLE OF DNA

Genetically the type of protein they go on the ladder down the middle to
modified food to make. For example, the produce two single strands. These
combination GGA is the codon for act as templates for the production
In agriculture, crops may be glycine. Sixty-four possible triplets of a second complementary
engineered to enhance them can be made from the four base DNA strand on each of them by
in some way. A genetically
pairs, and 61 of them code for a matching up the appropriate base
altered crop is known as a
genetically modified organism particular amino acid. The other pairs. The process results in two
(GMO). Companies that three act as signals such as “start” strands of whole DNA that are
operate in this sector may and ”stop,” which govern how exactly the same as the original.
modify a plant’s DNA so that information is read by the cellular Since DNA remains in the
it produces more of a certain machinery. DNA is also organized nucleus of the cell, a related molecule
nutrient or a toxin specific to into separate chromosomes, of called messenger ribonucleic acid
a particular insect pest. The which there are 23 pairs in the (mRNA) copies segments of DNA
DNA of a plant may also be human cell. coding sequence and carries the
altered to become resistant to information to the regions of the
a particular herbicide, so that Copying the code cell where new proteins are made.
use of the chemical kills only When cells divide, DNA needs to RNA is chemically related to DNA,
the weeds and not the crop. be copied. This is achieved by the but the thymine base (T) is
Some ecologists argue that splitting of base pairs, which cuts replaced by the base uracil (U),
there is a risk of genetically
which is less stable but requires
unmodified plants becoming
less energy to make. Stable living
contaminated by GMOs. They The structure of DNA organisms benefit from having
also point out that the long-
term effects of eating such DNA genomes, but RNA makes up
foods are as yet not properly genomes of some viruses, where
understood. Another concern stability can be less advantageous.
is that in the future large DNA is found in all living things
agrochemical companies could on Earth, from amoebae to insects,
control the world’s food supply to trees, tigers, and humans. Of
by patenting the GMOs that course, the sequence of base pairs
they produce, to the detriment varies, and this difference allows
of poorer nations. geneticists to trace relationships
between different species.
Good and bad errors

adenine thymine DNA is a highly stable molecule,
but sometimes mistakes, known
as mutations, occur. These can be
in the form of an error, duplication,
or omission in the order of the
cytosine guanine nucleotides A, C, G, and T. Mutation
can be spontaneous—the result
of errors that occur when the DNA
is copied—or may be induced
by external influences such as
exposure to radiation or cancer-
causing chemicals. Some mutations
New kinds of rice are being have no effect, but others may
developed through genetic A DNA molecule consists of a double
helix formed by two strands, made up change what the gene produces
modification. This may improve
the nutritional value of the crop of sugars and phosphates, linked by or inhibit the functioning of a
or its resistance to disease. paired base nucleotides: adenine and gene. This can lead to problems in
thymine or cytosine and guanine. the organism as a whole. Examples

DNA barcoding
The idea of DNA barcoding
was first raised in 2003 when
a team at the University of
Guelph, Canada, suggested
that it would be possible to
identify species by analyzing
a common section of their
DNA. Led by Dr. Paul Hebert,
researchers chose a region in
the gene known as cytochrome
c oxidase 1 (“CO1”), made up
of 648 base pairs. This region
is quick to analyze, but the
sequence is still long enough
to differentiate between and
within animal species.
Different gene segments can
be used for other forms of life.
The first part of the
barcoding system involves
of disorders caused by gene Mutated blood cells occur in cataloguing samples of known
mutations include cystic fibrosis sickle-cell disease—a genetic disorder species. The DNA is extracted
and sickle-cell disease. passed on when both parents carry
and organized into a sequence
the faulty gene. It can be painful and
Although many mutations are increases the risk of serious infections. of base pairs, a process known
harmful, occasionally a mutation as “sequencing.” The sequence
will confer an advantage on an is then stored in a computer
individual, enabling it to survive in parent. Additionally, with access database, so that when a DNA
its environment better than others to such data it is possible to sample from an unknown
of the same species. This type of screen embryos for known genetic species is sequenced and
mutation may end up being passed disorders before implantation in the entered into the database, the
on through the process of natural womb. By March 2018, the DNA of computer will match it with
existing records. The barcoding
selection. Over many generations, around 15,000 organisms had been
technique has proved useful
mutation is a mechanism for sequenced. Such information can for taxonomy, helping classify
diversification, survival of the help show how animals are related animals and plants.
fittest, and ultimately evolution. in the evolutionary line and how
they have diversified.
The human genome While the discovery of the
On April 14, 2003, scientists composition and structure of DNA
completed the lengthy task of has revolutionized the science of
mapping (sequencing) the entire heredity, it is worth noting that
human genome. Geneticists worked the regions of DNA used for coding
out the precise position of all the proteins account for just 2 percent With genetic
base pairs in a chain of some three of the entire human genome. engineering, we will
billion of the base nucleotides The nature of the other 98 percent be able … to improve
comprising an estimated 30,000 is not yet fully understood by the human race.
individual genes. This has allowed geneticists, but it is believed that at Stephen Hawking
geneticists to identify new genes least some of these regions involve
and the role they play in organisms. the regulation of the way genes are
Armed with this knowledge, expressed, or activated. It seems
an individual can find out if they that many more discoveries await
have inherited a faulty gene from a future geneticists. ■
38
GENES ARE
SELFISH
MOLECULES
THE SELFISH GENE
T
he concept of the “selfish for the bodily types and behaviors
IN CONTEXT gene” was popularized (phenotypic traits) that successfully
by British evolutionary promote their own propagation.
KEY FIGURE
biologist Richard Dawkins in his Supporters of the theory argue that
Richard Dawkins (1941–)
1976 book of that name. It states because heritable information is
BEFORE that evolution is fundamentally passed through the generations by
1963 British biologist William based upon the survival of different the genetic material of DNA, both
Donald Hamilton writes about forms of a particular gene at the natural selection and evolution
the “selfish interests” of the expense of others. The forms that are best considered from the
gene in The Evolution of survive are those that are responsible perspective of genes.
Altruistic Behaviur.
1966 American biologist
George C. Williams proposes Natural selection works toward the survival
in his book Adaptation and of the gene, not the individual.
Natural Selection that altruism
is a result of selection taking
place at the level of the gene.
AFTER
1982 Richard Dawkins argues Animals that warn
in The Extended Phenotype Male black widow
others of approaching
spiders mate
that the study of an organism predators sacrifice
even though the
should include analysis of themselves at the
females eat them
how its genes affect the expense of the
immediately after.
surrounding environment. wider group.
2002 Stephen Jay Gould

critiques Dawkins’ theory in
The Structure of Evolutionary
Theory, which revisits and Nonbreeding bees in bee
refines the ideas of classical colonies serve to help the
Darwinism. community survive.

See also: Evolution by natural selection 24–31 ■ The rules of heredity 32–33
■ The role of DNA 34–37 ■ Mutualisms 56–59
circumstances in which the gene

can achieve its own selfish goals by
fostering apparent altruism in the
organism. One example is kin
selection, the evolutionary strategy
that favours the reproductive
success of an individual organism’s
relatives, even at the cost of the
individual’s own reproduction
or survival.
An extreme example of Richard Dawkins
genetically based altruism is
A male black widow spider gingerly eusociality. Honey bees are a eusocial Richard Dawkins was born
approaches a huge female to mate. This species. They live in colonies which in Kenya to British parents.
genetically driven act will reproduce include breeding and non-breeding After the family returned to
his genes but will lead to his death.
individuals. By helping the colony the UK, he developed a strong
survive, the many thousands of interest in the natural world
Dawkins was strongly influenced non-breeding worker bees ensure and studied zoology at Oxford
by the work of William Donald the reproduction of the genes they University. While there, he
Hamilton on the nature of altruism have in common with the sole was tutored by Nobel Prize-
and closely examined the biology breeding individual, the queen. winner Niko Tinbergen, who
of selfishness and altruism in The Critics of Dawkins’ theory argue was a pioneer of animal
behavior studies. After a
Selfish Gene. He argued that that since individual genes do not
brief period at the University
organisms were simply vehicles control behaviour, they cannot be
of California at Berkeley,
that supported their genes, or said to be acting selfishly. Dawkins Dawkins returned to Oxford
“replicators.” Genes that help an has maintained that he never to lecture in zoology.
organism survive and reproduce meant to suggest that genes had Richard Dawkins is best
tend also to improve those genes’ their own conscious will. He later known for his book The Selfish
own chances of being replicated. wrote that “the immortal gene” Gene, in which he argues
Successful genes often provide might have been a better title for that the gene is the principle
a benefit to the host organism. For both his concept and the book. ■ unit of selection in evolution.
example, a gene that protects an His theory later triggered a
animal or plant against disease series of fierce debates with
thereby helps that particular gene Stephen Jay Gould and other
to spread. However, the interests of evolutionary biologists.
the replicator and the vehicle may Dawkins is also known as a
sometimes seem to be in conflict. strong advocate of atheism
The theory of evolution and feminism.
Genes drive the male black widow
spider to mate despite the risk of is about as much
open to doubt as the Key works
being eaten by her. However, the
male’s sacrifice nourishes the theory that the Earth
1976 The Selfish Gene
female and improves the prospect goes around the Sun. 1982 The Extended Phenotype
of his genes being passed on. Richard Dawkins 1986 The Blind Watchmaker
2006 The God Delusion
Selfishness and altruism 2009 The Greatest Show
Gene selfishness usually gives rise on Earth: The Evidence
to selfishness in the behavior of an for Evolution
individual organism, but there are
ECOLOGI
PROCESS
CAL
ES
42 INTRODUCTION
Robert MacArthur’s Dan Janzen observes the

Joseph Grinnell publishes research on North American interdependence of acacia
his research on the California warblers shows how different trees and the ants that reside
Thrasher, establishing the species can avoid directly on them, and concludes that
basis for the theory of competing with each other the species evolved in a
ecological niches. in order to coexist. mutualistic manner.
1917 1957 1965
1925–26 1961 1969
The Lotka-Volterra model uses Joseph Connell reveals that Robert Paine
a mathematical equation to different types of barnacle coins the term “keystone
describe the interactions between thrive in different tidal species” to describe species
predator and prey. zones, although they could, that play a crucial role in
in theory, live in any of them. ecosystem functions.
I
n the 5th century BCE, the Greek beyond their own local area. As Alfred Lotka introduced one of the
historian Herodotus described technology improved and people first mathematical models ever
watching crocodiles open their began to travel the world, scientists applied to ecology. Now known
jaws for plovers to pick food from such as Robert Hooke, Antonie van as the Lotka-Volterra model, its
their teeth. He may have been the Leeuwenhoek, Carl Linnaeus, predator–prey equations help
first to write about an ecological Alexander von Humboldt, Alfred predict the population fluctuations
process—in this case a mutualistic Russel Wallace, Charles Darwin, of these two groups.
relationship between reptiles and and Johannes Warming became In the early years of the 20th
birds. Aristotle and Theophrastus increasingly aware of ecological century, Joseph Grinnell conducted
observed many more interactions processes and laid the foundations extensive research into animals’
between animals and their of the science of ecology, even if habitat needs in the western United
environment in the 4th century BCE. they didn’t use that word. States. He observed that species
Over the next two millennia, had different “niches” within a
countless other observations of the Mathematical models habitat—and that if two species
natural world were made, but a deep It had long been understood that have approximately the same food
understanding of how organisms one of the most basic ecological requirements, one will “crowd out”
interacted with each other and the processes is the struggle for the other. Darwin had observed this
world around them was hampered survival: for herbivores to find food, on his travels aboard HMS Beagle,
by the inability to observe very predators to find prey, and prey to but Grinnell’s axiom developed the
small things, those that were active avoid being eaten. Predators do idea further, as did subsequent
at night, or those living underwater. everything they can to hunt and research. In 1934, Georgy Gause
Additionally, few people with an eat prey, and the latter do all they demonstrated what he called the
interest in nature experienced much can to avoid being eaten. In 1910, competitive exclusion principle in
ECOLOGICAL PROCESSES 43
Research published by
Ronald Pulliam, Eric Charnov, Robert Sterner and James Elser
and Graham Pyke expands on pioneer the study of ecological
Roy Anderson and Robert the optimal foraging theory that stoichiometry—how ratios of
May demonstrate how animals try to gather resources different chemicals within
epidemics affect animal while wasting as little living organisms change
population growth rates. energy as possible. with certain reactions.
1970s 1977 2002
1972 1991
Knut Schmidt-Nielsen Earl Werner publishes

publishes How Animals his findings about
Work. The book hugely nonconsumptive
influences the field effects of predators
of ecophysiology. on prey.
laboratory projects. As William E. to play, too—as Earl Werner New technology

Odum put it in 1959, “the ecological demonstrated 30 years later. His Technological advances—including
niche of an organism depends not work revealed the non-consumptive sophisticated chemical sampling
only on where it lives, but also on impact of predatory dragonfly larvae techniques, satellites with remote
what it does.” on the behavior and physical sensing equipment, and computers
development of their tadpole prey. capable of rapidly processing huge
From field to lab Since the mid-20th century, quantities of data—have opened
Laboratory experiments and field many new ideas on ecological up new areas of study.
observations are the main methods processes have emerged. Work by Ecological stoichiometry, for
of providing data for the study Robert MacArthur and others on example, studies the flow of energy
of ecological processes, but field competition between species led to and chemical elements throughout
experiments—in which a local the development of optimal foraging food webs and ecosystems, from the
environment is manipulated to test theory, which seeks to explain why molecular level up. Like so many
a hypothesis—were not conducted animals choose to eat some food ideas in ecology, its origins can be
with scientific rigor until Joe items and not others. Mutualistic traced back many years, but only
Connell’s work with barnacles relationships became better took hold with Robert Sterner and
in Scotland. His experiments—the understood through the research James Elser’s 2003 book Ecological
results of which were published of biologists such as Daniel Janzen. stoichiometry: The biology of
in 1961—were meticulously planned Robert Paine’s work with starfish elements from molecules to the
and observed, and were repeatable. and mussels also highlighted the biosphere. New techniques such
Connell set the “gold standard” concept of keystone species— as this will undoubtedly continue
for fieldwork, but experiments in those that have a disproportionate to deepen our understanding of
laboratories still have a vital role influence on their ecosystems. processes in ecology. ■
LESSONS FROM
MATHEMATICAL THEORY
ON THE STRUGGLE
FOR LIFE
PREDATOR–PREY EQUATIONS
46 PREDATOR–PREY EQUATIONS
T
he predator–prey equations
IN CONTEXT Populations of two species, are an early example of the
one predator, the application of mathematics
KEY FIGURES
other prey, interact. to biology. Formulated in the 1920s
Alfred J. Lotka (1880–1949),
by American mathematician Alfred
Vito Volterra (1860–1940) J. Lotka and Italian mathematician
BEFORE and physicist Vito Volterra, the
1798 British economist two equations—also known as
Thomas Malthus shows that The prey has access to the Lotka–Volterra equations—
the rate at which the population food and its population describe the way in which the
growth is exponential. population of a predator species
changes increases as the size
of the population grows. and that of its prey fluctuate in
relation to each other.
1871 In Lewis Carroll’s novel Lotka proposed the equations
Through the Looking Glass, in 1910, as a way of understanding
the Red Queen tells Alice, When prey animals meet the rates of autocatalytic chemical
“you have to run just to stay a predator, they reactions—chemical processes
in the same place.” are eaten. that regulate themselves. In the
following decade, he applied
AFTER the equations to the population
1973 American biologist Leigh dynamics of wild animals.
Van Valen proposes the Red In 1926, Vito Volterra arrived
Queen effect, which describes Eating prey results at the same conclusions. He had
the constant “arms race” in more predators. become interested in the subject
between predators and prey. after meeting Italian marine
biologist Umberto D’Ancona.
1989 The Arditi–Ginzburg D’Ancona told Volterra how the
equations offer another model percentage of predatory fish
of predator–prey dynamics caught in nets in the Adriatic
More predators
by including the impact of the Sea had greatly increased during
results in less
ratio between predator and prey. World War I. This change was
prey, reducing the
number of predators. clearly linked to the drastic
reduction in fishing during the
Vito Volterra Born in 1860 in Ancona, Italy, the Volterra refused to swear loyalty
son of a Jewish cloth merchant, to Italy’s fascist dictator Benito
Vito Volterra grew up in poverty. Mussolini and was dismissed
Despite this, in 1883, aged just 23, from the University of Rome.
he secured a position as professor Forced to work abroad, he only
of mechanics at the University returned to Italy for a short time
of Pisa and began a career as before his death in 1940.
a mathematician. Further
professorships at the universities Key works
of Turin and Rome followed. In
1900, Volterra married, fathering 1926 “Fluctuations in the
six children, although only four Abundance of a Species
survived to adulthood. He was Considered Mathematically,”
made a senator of the Kingdom Nature
of Italy in 1905 and worked on the 1935 Les associations
development of military airships biologiques au point de vue
during World War I. In 1931, mathématique
See also: Evolution by natural selection 24–31 ■ The selfish gene 38–39 ■ Ecological niches 50–51 ■ Competitive exclusion
principle 52–53 ■ Mutualisms 56–59 ■ Keystone species 60–65 ■ Optimal foraging theory 66–67
A cheetah pursues a Thomson’s increases as the population grows. occurring factors. As a result, wild
gazelle. The predator–prey equations From this theory, Malthus predicted populations should in theory be
are able to model the way populations a catastrophic future for humanity. more or less static, fluctuating only
of both species will change in response
to the activities of the other.
The number of humans was around the carrying capacity,
growing much more quickly than assuming the random impacts of
the amount of food that could be catastrophic events are ignored.
war years, but D’Ancona could not produced by the world’s farmlands. However, this relative
explain why less fishing did not Eventually, Malthus argued, a point equilibrium did not always match
produce more fish of all kinds in the would be reached when the human up with observations—as in ❯❯
nets. Using the same equations as population would succumb to
Lotka, Volterra eventually explained global famine and decline.
the fluctuations in both the predator Malthus’s bleak vision did not
and the prey species. happen, thanks to technological
advances in agriculture and the
Population principles development of artificial fertilizers,
At the time Lotka and Volterra but his population model became The food species
made their calculations, the science applicable to species populations cannot, therefore, be
of population dynamics was still within ecosystems. Every habitat, exterminated by the
in its infancy, having barely moved and the niche occupied by a species predatory species, under
on since the population studies of within its community of organisms, the conditions to which
British economist Thomas Malthus has a carrying capacity—the our equations refer.
in the late 18th century. According maximum population that can Alfred J. Lotka
to Malthus’s theory, a population be supported by the resources
grows or declines rapidly as long available, such as water, space,
as the environmental factors for food, and light. Any rise in
survival are constant, and the rate population above this level is
at which that population changes likely to be reduced by naturally
48 PREDATOR–PREY EQUATIONS
no reproduction limits and the species and the predation rate.
rate of change in a population For example, oscillations in the
is proportional to its size; second, size of an ant population and that
that the prey population—presumed of an anteater are barely noticeable
to be a herbivore—is always able because they reproduce at such
Mathematics without to find enough food to survive. different rates. The oscillations
natural history is sterile, but Next, they assumed that the prey in the populations of species that
natural history without population is the predators’ only breed at similar rates, such as the
mathematics is muddled. source of nourishment, and that Iberian lynx and rabbit, are much
John Maynard Smith the predators never become full more pronounced.
British mathematician and never stop hunting. Finally,
and evolutionist they assumed that environmental Nature’s arms race
conditions, such as weather or The predator–prey equations
natural disasters, had no impact revealed that species are locked
on the process. The effect of the together in a never-ending struggle,
genetic diversity of the predators swinging from near disaster and
and prey animals on their ability to extinction to times of abundance
survive was not taken into account. and fertility. In this biological “arms
D’Ancona’s account of a sudden When plotted on a graph, the race,” the evolutionary pressure
increase in the population of predator population trails the rise on the prey species is to escape
predatory sea fish. One theory and fall of the prey population, and predation and survive, so as to have
to explain this discrepancy is still rising as the prey population more offspring. Meanwhile, the
started from the premise that the starts to decline. This explained predator is under pressure to have
population of predators is related D’Ancona’s observation of the larger a higher predation rate in order
to the size of the population of their proportion of predators after the to provide food for more offspring.
food supply, such as prey species. prey population had been allowed However, neither species is
The relationship suggests that to boom by a reduction in fishing. superior, responding instead
when a lot of food is available, there The relative fluctuations of the to the adaptations of the other. The
will be a large predator population. populations depends on the relative predator–prey relationship between
The growing predator population reproductive rates of the two even-toed hoofed mammals—such
should then begin to reduce the
amount of prey, which will in
turn lead to a drop in the number Predator–prey population cycles
of predators. The size of both
KEY The predator and prey
populations will rise and fall, but populations rise and fall
the ratio of predators to prey will Prey over time in regular cycles.
remain stable. Predator Although the degree to
Such a balanced theory was still which they change varies,
at odds with species observations. the cycle follows a broadly
Through mathematical modeling, similar pattern.
POPULATION
Volterra was able to show that the

average sizes of predator and prey
populations do indeed oscillate but
the rate at which each population
is growing or declining is always
changing and almost never
matches the changes experienced
by the other population. To
eliminate variables, Volterra made
a series of assumptions: first, that
the prey and predator species have TIME
as antelopes and deer—and
mammalian carnivores, like the
big cats and wolves, is an example
of this evolutionary arms race. The
hoofed animals have long legs,
extended by walking on the very
tips of thickened and fused toe
bones. This adaptation allows
them to outrun and outjump their
predators. In response, big cats—
such as lions and tigers—have
evolved speed and strength to
bring down large, fleet-footed prey
in surprise attacks. Wolves have
evolved the stamina to run for
long distances without stopping.
This allows them to work as a
team to chase down their prey
and kill them when the exhausted
prey collapse.
While the predator–prey
equations offer an insight into
the population dynamics of two
species, the assumptions they rely
on are rarely reflected in real life.
Some predators do specialize in
killing a single prey species, but
other factors in the ecosystem
also affect their populations.
Other applications
The Lotka–Volterra equations have
been used to study the dynamics of another species but also the prey The parasitoid wasp lays its eggs
food chains and food webs in which species of a third. They have in aphids (the smaller, yellow insects
one species may be a predator of also been used to examine the shown above). It is called a parasitoid
because the wasp’s larvae later eat the
relationship between host and aphids as they grow.
parasite species, which bears
some resemblance to that between
prey and predator. Parasites often thanks to beneficial genes, certain
specialize in one host species— individuals in a host population
Volterra was interested a relationship that should resemble are able to maintain their fitness
in a mathematical the one described by the Lotka– despite the attacks from parasites.
theory of ‘the survival Volterra equations. However, in The parasites constantly evolve to
practice the process of evolution exploit these seemingly immune
of the fittest.’
is thought to interfere with this. individuals, and therefore the
Alexander Weinstein A parasite does not usually kill beneficial genes in the host
Russian mathematician
its host (those that do are called population also change. In this way,
parasitoids), but can reduce its evolution is happening all the time,
fitness. The Red Queen evolutionary as the parasite and host battle it
theory, proposed in the 1970s by out—although everything appears
Leigh Van Valen, describes how, to stay the same. ■
50
EXISTENCE IS DETERMINED
BY A SLENDER THREAD
OF CIRCUMSTANCES
ECOLOGICAL NICHES
A
n organism’s niche is competition with other species.
IN CONTEXT a combination of its For ecologists, a full knowledge
place and its role in the of an organism’s niche is vital to
KEY FIGURE
environment. It encompasses how inform interventions to compensate
Joseph Grinnell (1877–1939)
the organism meets its needs for for the environmental changes
BEFORE food and shelter, as well as how it caused by habitat destruction and
1910 In a paper about beetles, avoids predators, competes with climate change.
Roswell Hill Johnson, a US other species, and reproduces. The pioneer of the niche
biologist, is the first person All its interactions with other concept was Joseph Grinnell, a US
to use the word “niche” in organisms and the nonliving biologist who studied a bird called
a biological context. environment are also part of what the California Thrasher. In 1917, he
makes up its niche. A unique niche published his observations, which
AFTER is an advantage for any animal or showed how the bird fed and bred
1927 British ecologist Charles plant because this reduces in the underbrush of a scrubby
Elton stresses the importance
of an organism’s role as well as
its “address” in his definition of
an ecological niche in his book There is constant
Animal Ecology. competition for food and
resources; better adapted Reducing competition
1957 In an academic paper species outcompete those increases the chances
called “Concluding Remarks,” less suited to the of survival.
British ecologist George environment.
Evelyn Hutchinson expands
the theory of niches to
embrace an organism’s entire
environment.
1968 A study by Australian Existence of each
D.R. Klein of the introduction, species is Finding a unique niche
increase, and die-off of reindeer determined by a is the circumstance that
on St. Matthew Island, Alaska, slender thread removes competition.
identifies the destructive niche. of circumstances.
See also: Competitive exclusion principle 52–53 ■ Field experiments 54–55 ■ Optimal foraging theory 66–67
■ Animal ecology 106–113 ■ Niche construction 188–189
An ultra-specialist Giant pandas occupy a very Pandas eat different parts of

specialized ecological niche, as bamboo plants according to the
their diet consists mainly of seasons. In late spring, they
bamboo. Bamboo is a poor food prefer the first green shoots.
source, low in protein and high in They eat leaves at other times
cellulose. Pandas can digest only of the year, and stems in winter
a small proportion of what they when little else is available.
eat, which means they have to Pandas have evolved muscular
eat a lot of bamboo—as much as jaws and a pseudothumb to
28 lb (12.5 kg) each day—and manipulate bamboo stems. Their
forage for up to 14 hours a day. digestive tract is inefficient at
It is unclear why pandas have processing large quantities of
become so dependent on bamboo, plant material because it
but some zoologists suggest it is remains similar to that of its
because it is an abundant and carnivorous ancestors, although
reliable food source, and pandas digestion is helped by the
are not skilled predators. bacterial fauna in their gut.
habitat known as chaparral, and factors. Thirty years later, George coexist (niche partitioning), and
how it escaped predators by Evelyn Hutchinson expanded the the overlap of resources by different
running through the underbrush. definition yet further. He argued animals and plants (niche overlap).
The thrasher’s camouflage, short that a niche should take into
wings, and strong legs were account all of an organism’s The importance of habitat
perfectly adapted for life in this interactions with other organisms Ecological niches depend on the
environment. Grinnell saw the and its nonliving environment, existence of a stable habitat; small
chaparral habitat as the thrasher’s including geology, acidity of soil or changes can eradicate niches that
“niche.” His idea also allowed for water, nutrient flows, and climate. organisms once filled. For example,
“ecological equivalence” in plants Hutchinson’s work encouraged dragonfly larvae only develop
and animals, whereby species others to explain the variety of within a certain range of water
distantly related and living far resources used by a single acidity, chemical composition,
apart could show similar organism (niche breadth), the ways temperature, and prey, and with
adaptations, such as feeding in which competing species a limited number of predators.
habits, in similar niches. In the The right vegetation is needed by
Australian outback, for instance, adult females for egg-laying, and
babbler bird species forage in the by larvae for metamorphosis.
scrubby vegetation in a similar way The dragonfly also impacts its
to the unrelated thrasher. Grinnell environment: its eggs are food for
also identified “vacant” niches— amphibians; its larvae, which are
habitats that a species could [A niche] is a highly both predators and prey, add
potentially occupy, but where it abstract multi- nutrients to the water; and the
was not present. dimensional hyperspace. adults prey on insects. These
George Evelyn requirements and impacts define
Widening the niche Hutchinson its ecological niche. Hutchinson
In the 1920s, ecologist Charles argued that for a species to persist,
Elton looked beyond a simple conditions had to be within the
habitat definition for “niche.” For required ranges. If conditions moved
him, what an animal ate and what outside the niche requirements, a
it was eaten by were the primary species could face extinction. ■
52
COMPLETE
COMPETITORS
CANNOT COEXIST
COMPETITIVE EXCLUSION PRINCIPLE
C
ompetition is the driver of adapts, so that it no longer
IN CONTEXT evolution; the need to be competes. This proposition, known
bigger, stronger, and better as the “competitive exclusion
KEY FIGURE
inevitably leads to adaptations that principle,” was set out by Russian
Georgy Gause (1910–86)
give a species an edge. When two microbiologist Georgy Gause and
BEFORE species compete for identical is also known as Gause’s Law.
1925 Alfred James Lotka first resources, the one which has any Gause devised his principle
uses equations to analyze advantage will outdo the other. As from laboratory experiments, using
variations in predator–prey a result, the weaker of the two cultures of microorganisms, rather
populations, as does species either becomes extinct or than from observations in nature. In
mathematician Vito Volterra,
independently, a year later. How warblers coexist
1927 Volterra enlarges and
updates his 1926 study to
include various ecological
interactions within
communities.
AFTER
1959 G. Evelyn Hutchinson
extends Gause’s ideas and Cape May Blackburnian Black-throated
produces a ratio describing the Warbler Warbler Green Warbler
limit of similarity between two
competing species.
1967 Robert MacArthur and
Richard Levins use probability Five species of
theory and Lotka–Volterra warblers are able to
equations to describe how share the same tree,
coexisting species interact. because each inhabits
its own “niche.” Living
in this way, without
Bay-breasted Yellow-rumped much overlap, the
Warbler Warbler birds do not compete.
See also: Evolution by natural selection 24–31 ■ Ecological niches 50–51 ■ Animal ecology 106–113 ■ The ecosystem
134–137 ■ The ecological guild 176–177 ■ Niche construction 188–189 ■ Invasive species 270–273
genetics. In fact, the competitive

exclusion principle—although a
useful theoretical model—is rarely
seen in nature, simply because, in a
bid to survive, a weaker competitor
tends to quickly move on or adapt. Let us make for this purpose
an artificial microcosm… let
Avoiding competition us fill a test-tube with nutritive
Most creatures can make the medium and introduce to it
changes necessary for survival. A several species of protozoa
variety of birds can live in a garden consuming the same food
during any one year because they or devouring each other.
all operate in different “niches.” Georgy Gause
They have contrasting beak shapes
and sizes that allow them to eat
The red squirrel is smaller than the
gray, and has a more restricted diet different types of food—the robin
and habitat. Reds may also die from the preferring insects, the finch eating
squirrel parapoxvirus, which is carried seeds. Their choice of habitat and
by the grays but does not affect them. feeding times might also vary; this
is known as resource partitioning. at different heights and depths of
In 1957, Robert MacArthur the foliage. In this way they avoid
nature, he proposed, there were too noted this phenomenon in North competing with each other.
many variables to draw conclusions American warblers. The five
about how ecological mechanisms species he observed, each with An invasive competitor
work. He argued that little progress distinctive, colorful markings, Problems often arise if an exotic
had been made since Darwin’s era flitted in and out of coniferous species is suddenly introduced
in understanding how species trees, feeding on bugs and other to an ecosystem. Britain’s red
compete for survival, whereas the insects. They could coexist in one and gray squirrels provide a clear
experimental method had produced habitat because they did not try to example. When the grey arrived
great advances in areas such as feed in the same part of the tree but from America in the 1870s, both
squirrel species competed for the
Types of competition same food and habitat, which put
the native red squirrel populations
The Competitive Exclusion “limiting resource,” the one that under pressure. The gray had
Principle covers two main types both require in order to breed. the edge because it can adapt
of competition. Intraspecific Ecologists make a further two its diet; it is able, for instance, to
competition is between distinctions. Interference is eat green acorns, while the red can
individuals of the same species when two organisms fight only digest mature acorns. Within
and ensures the survival of directly with each other over a the same area of forest, gray
the fittest, so that only the limited resource, such as a mate squirrels can decimate the food
healthiest individuals—or those or a preferred food. Exploitation supply before red squirrels even
best adapted to a particular is indirect competiton, such as have a nibble. Grays can also live
environment—will breed. The stripping out a resource so there more densely and in varied
second type is interspecific: is none left for the competitor;
habitats, so have survived more
competition between two this can be seen in plants, when
different species that rely on a species’ uptake of nutrients or easily when woodland has been
the same resources. The most water is more efficient than that destroyed. As a result, the red
important of these will be the of its neighbors. squirrel has come close to
extinction in England. ■
54
POOR FIELD
EXPERIMENTS
CAN BE WORSE
THAN USELESS
FIELD EXPERIMENTS
E
xperimentation is crucial have been recognized. Before
IN CONTEXT in ecology. Without it, our the 1960s, experiments outside
ideas about why organisms a laboratory were a rarity.
KEY FIGURE
behave the way they do would be A laboratory, however, is an
Joseph Connell (1923–)
largely speculative. Rigorous artificial environment, where
BEFORE observation is also essential, but, organisms may not behave as they
1856 British scientists John much of the time, experimentation do in their natural habitat. For
Lawes and Joseph Gilbert start is needed for a full understanding example, bats leaving a roost at
the Park Grass Experiment at of those observations. dusk may follow different routes
Rothamsted, to test how Three main types of ecological to their foraging areas in spring
different fertilizers affect the experiments are used to test and late summer. The potential
yield of hay meadows. theories: mathematical models, reasons for the switch—changes
laboratory experiments, and field in prey distribution and predator
1938 Harry Hatton, a French experiments. Each method has its threats; seasonal differences in
ecologist, conducts one of merits, but it is only recently that tree cover; or human disturbance
the first marine ecology field the benefits of field experiments and light pollution—cannot
experiments, on barnacles be established in a laboratory.
on the Brittany coast. Mathematical modeling might help
predict patterns, but would be less
AFTER effective at identifying the causes
1966 American ecologist of change. To understand the bats’
Robert Paine removes the behavior, a study of their natural
starfish Pisaster ochraceus from environment is crucial, and
tide pools in a Pacific coast this is achieved only through
ecosystem, to test the effect research in the field.
of its absence on other species. Field experiments allow different
factors to be manipulated to test
1968 The Experimental Lakes their relevance. In the bat example,
Area, comprising 58 freshwater
lakes, is established in Ontario,
Canada, to study the effects Rain forest ecosystems are some of
the most species-rich environments
of nutrient enrichment
on Earth. This makes them especially
(eutrophication). valuable sites for ecologists to conduct
experiments in the field.
See also: Ecological niches 50–51 ■ Modern view of diversity 90–91 ■ Animal behavior 116–117 ■ The ecosystem 134–137
■ Niche construction 188–189
street lights could be switched off to Joseph Connell’s barnacle experiment

evaluate the impact of light pollution
on their behavior change. Chthamalus Highly desiccated
area during low tide
Balanus
Scottish barnacles High tide
In 1961, American ecologist Joseph
Connell published the results of
his research on barnacles on the
Scottish coast. Since free-swimming
barnacle larvae can settle anywhere,
Connell had tested why the lower
part of the intertidal zone was This experiment showed Fundamental
colonized by Balanus balanoides that Balanus could live only niches
barnacles and the upper part by in the lower intertidal zone, Realized
Chthamalus stellatus. He wanted to while Chthamalus could live niches
know if this was due to competition, in both the upper and lower
predation, or environmental factors. zones, but was outcompeted
by Balanus in the lower zone.
Connell manipulated the local
environment, and monitored it for
over a year. In one area, he removed Ocean
the Chthamalus barnacles. They
were not replaced by Balanus,
which suggested that Balanus could Low tide
not tolerate the desiccation that
occurred in the upper zone at low
tide. Connell then removed the species could live in the lower zone, successful when their nearest
Balanus population from the lower but only one could survive higher neighbor was of the same species.
zone, and found that Chthamalus up. This suggested that Chthamalus Each species is targeted by specific
barnacles did replace them. Both was better able to deal with the herbivores and pathogens, which
harsh conditions of the upper zone, will also eat or attack smaller,
but was outcompeted by Balanus weaker individuals of the species
lower down. The “fundamental nearby. This prevents “clumping”
niche” of Chthamalus (where the of one tree species.
species would normally be able to In 1978, Connell proposed
[Connell’s] studies … have survive) encompassed both zones, the intermediate disturbance
improved our understanding but its “realized niche” (the actual hypothesis (IDH). This states
of the mechanisms area it inhabits) was more restricted. that both high and low levels of
that shape population disturbance reduce species
and community Diversity experiments diversity in an ecosystem, so the
In the early 1970s, Connell and greatest range of species can be
dynamics, diversity,
American ecologist Daniel Janzen expected between those extremes.
and demography. published an explanation of the Several studies support IDH. One,
Stephen Schroeter degree of tree diversity in tropical carried out in waters off Western
Marine scientist
forests: the Janzen–Connell Australia, examined the effects of
hypothesis. Connell mapped wave disturbance on diversity.
trees in two rain forests in North Species diversity was found to be
Queensland, Australia, and found low both at exposed offshore sites
that seedlings tended to be less and at sheltered sites. ■
56
IN CONTEXT
MORE NECTAR KEY FIGURE

Dan Janzen (1939–)
MEANS MORE ANTS

BEFORE
1862 Charles Darwin proposes
that an African orchid with
AND MORE ANTS

a long nectar receptacle must
be pollinated by a moth with
MEAN MORE NECTAR

an equally long proboscis.
1873 Belgian zoologist Pierre-
Joseph van Beneden first
MUTUALISMS uses the term “mutualism”
in a biological context.
1964 The term “coevolution”
is first used by American
biologists Paul Ehrlich and
Peter Raven to describe
the mutualistic relations
between butterflies and
their food plants.
AFTER
2014 Researchers discover
an unusual yet beneficial
three-way mutualism involving
sloths, algae, and moths.
I
n biology, there are several
kinds of interaction between
organisms. One species in an
ecosystem may lose out to another
when competing for the same
resources. A prey species may be
eaten by a predator. There are also
symbiotic relationships, in which
one species benefits but not at the
expense of the other, or where one
organism does not benefit but still
survives. In the relationship known
as “mutualism,” both organisms
benefit from the relationship.
A tree and its ants

In the mid-1960s, Daniel Janzen,
a young American ecologist,
became fascinated by the amazing
See also: Evolution by natural selection 24–31 ■ Ecological niches 50–51
■ Competitive exclusion principle 52–53 ■ Animal ecology 106–113
Yuccas and
their moths
In the hot, arid regions
of the Americas, there is
a remarkable mutualistic
relationship between yucca
shrubs and yucca moths. No
other insects pollinate these
plants, and no other plants
host yucca moth caterpillars.
A female yucca moth collects
pollen from the flower of one
yucca plant and deposits it in
the flower of another yucca,
fertilizing the plant as it does
so. The moth then cuts a hole
in the flower’s ovary and lays
an egg; she may lay several
in the same flower. When the
eggs hatch, the caterpillars
feed on the seeds developing
in the flower but do not eat
them all, leaving enough for
the plant to propagate. If too
many eggs are laid in one
mutualistic relationship between Ants and their larvae shelter inside flower, the plant sheds it
acacia trees and ants in eastern the swollen thorn of an East African before the caterpillars hatch –
whistling thorn acacia tree. In return
Mexico. His research was one of leaving those insects to
the ants swarm from their nests
the first in-depth studies of such to protect the tree from herbivores. starve. Without these moths,
an interaction. The two partners the yuccas would not pollinate
were the swollen-thorn acacia and and would soon die out.
the acacia ant, which lives in the Without the ants, a tree would be Without the yuccas, the moths
bullhorn-shaped thorns of the tree. stripped of its leaves and die within would have nowhere to lay
He found that queen ants sought six months or a year. Because it and nurture their eggs, and
out unoccupied shoots, cut a hole could not sustain growth, it was they too would not survive.
in one of the swollen thorns, and also likely to be shaded out by
laid their eggs, sometimes leaving competing trees. Janzen clipped
the thorn to forage on the tree’s thorns and cut or burned shoots
nectar. Larvae hatching from the to remove ants from trees, and
eggs then fed on the acacia’s leaf- found that the ants moved back in
tips, with their rich supplies of when new thorns started to grow.
sugars and proteins. The larvae In return for food and shelter,
later metamorphosed into worker the ants provided two services
ants. In time, all the tree’s thorns for the tree: they defended its
became occupied, with up to foliage from leaf-eating insects
30,000 ants living in a colony. and ate potentially competitive tree
Janzen showed that, unless the seedlings growing close by. Janzen
acacia ants were present to defend described the acacias and their
it, the swollen-thorn acacia lost ants as “obligate mutualists”,
the ability to withstand damage meaning that one species would
caused by insects that ate its die out without the other. If the ants
leaves, stems, flowers, and roots. were removed, the swollen-thorn ❯❯
58 MUTUALISMS
1. Adult female wasp

There is
mutual aid
Life cycle of the in many species.
fig wasp Pierre-Joseph
van Beneden
Belgian zoologist
4 . New-generation
2. Wasp enters fig, adult female wasps
lays eggs, pollinates pick up pollen and
flowers, and dies emerge from fig
and thence disperse their spores.

3. Inside mature fig,
When a bird swallows a fruit, it
male wasps fertilize carries the seeds with it
new females and dig as it flies away; the indigestible
escape tunnels for them seeds may be excreted in faeces
far from where they were eaten.
The fig wasp and the fig share a complex In all these situations, the plants
service-resource mutualism, in which the wasp provides provide a resource (food) and the
the service of pollination and the fig plants provide mammals, flies, and birds provide
a secure environment for the wasp eggs to develop. a service (transport).
However, not all mutualistic
acacia would have no means of the animal. It is estimated that relationships involve plants. In
defending itself. And if the acacia nearly three-quarters of flowering Africa, birds named oxpeckers
trees were removed, the ants would plants (some 170,000 species) are and herbivorous mammals such
have no home. pollinated by 200,000 animal as impalas and zebras practise
species. Typically, a pollinating another kind of service-resource
Benefits for all insect is attracted to a flower by its mutualism. The oxpeckers pick
There are two fundamental types colours or scent to drink nectar or ticks from the mammals’ fur,
of mutualism—service-resource collect pollen, and pollen attaches removing irritation and a source
and service-service relationships. to part of the insect’s body to be of disease, while at the same time
They are defined by the nature carried to the next flower, where having a meal. Oxpeckers also
of the relationship between the it is deposited. The flower and its make loud calls when they sense
partner organisms, whether it is pollinator have evolved to make danger, alerting the mammal host
the provision of a service or the this mechanism work effectively. as well as other oxpeckers.
supply of a resource—both are Some plants have also evolved In the insect world, some ants
usually key to survival. Service- a service-resource relationship in and aphids carry out a different
resource relationships are common which birds and mammals disperse form of service-resource mutualism.
in nature, with the fertilization, or their seeds, spores, or fruit. Seeds While the aphids feed on plants,
pollination, of flowers by butterflies, may become attached to the fur of the ants protect the aphids.
moths, bees, flies, wasps, beetles, a mammal browsing the plant’s Subsequently, the ants consume
bats, or birds the most widespread leaves; when the mammal wanders the honeydew that the aphids
example. The resource (pollen) is away, it disperses the seed. The release, using a “milking” process
provided by the flower, and the vile odor of stinkhorn fungi attracts on their smaller partners, by
service (pollination) is provided by flies, which lick the fungi’s slime stroking them with their antennae.
Service-service mutualisms, in return for the protection offered of orchids. Like many other
which both organisms offer each by the sea anemones’ venomous flowering plants, orchids rely on
other protection, are far less tentacles, the clownfish deters insects to pollinate them. Some
common. One unusual relationship predatory butterfly fish, removes have extraordinary structures in
takes place in the western Pacific parasites from its host, and also which to hold nectar and pollen.
Ocean, between around 30 species provides nutrients from its faeces. To lure the insect pollinators, the
of clownfish and 10 species of plants offer them a drink of energy-
venomous sea anemones. The sea Cooperative evolution giving nectar. This fascinated
anemones’ stinging, toxin-filled Relationships between service and Darwin, who was given a specimen
nematocysts, or capsules, on their resource providers have developed of the Madagascar orchid in 1862.
tentacles kill most small fish that over millions of years in a process The flower stores its nectar in a
come close, but not the clownfish. called “coevolution”—the evolution hollow spur nearly 30 cm (12 in)
Its thick layer of protective mucus of two or more species that affect long. Darwin and Wallace
provides immunity against the each other reciprocally. speculated that only a large moth
anemone’s sting, allowing the fish The term coevolution was could have a proboscis long enough
to live within the tentacles. In coined by American biologists Paul to reach the nectar—a theory
Ehrlich and Peter Raven in 1964, eventually proven in 1997. If the
but a century before the word orchid’s spur were shorter, a moth
The clownfish and sea anemone
could both survive without the other’s existed, the naturalists Charles could drink without picking up
protection, but their coevolved mutual Darwin and Alfred Russel Wallace pollen and so would not pollinate
relationship gives them a much higher were already aware of the concept, the flower. If the spur were longer,
chance of survival. not least through their observation a moth would not visit. ■
WHELKS
ARE LIKE LITTLE
WOLVES
IN SLOW MOTION
KEYSTONE SPECIES
62 KEYSTONE SPECIES
A
keystone species plays
IN CONTEXT a crucial role in the way
an ecosystem functions,
KEY FIGURE
even though it is often a small
Robert Paine (1933–2016)
part of the overall biomass of the
BEFORE ecosystem. Because it exerts a Do you want an auto
1950s In Kenya, farmer and disproportionately large effect on mechanic who…can name,
conservationist David the environment relative to its list, and count all of the parts
Sheldrick introduces elephants biomass, if a keystone species of your engine, or one who
to Tsavo East National Park, disappears from an ecosystem, really understands how each
and discovers this results in a that ecosystem will change part interacts with the others
major increase in biodiversity. dramatically. The importance of to make a working engine?
keystone species was brought to Robert Paine
1961 Fieldwork by American light by the American biologist
ecologist Joseph Connell on Robert Paine—who derived the
Scotland’s rocky shores shows term from the central “keystone” at
that removing predatory the top of an arch that stops it from
whelks alters the distribution collapsing—in his 1969 article “A
of their barnacle prey. Note on Trophic Complexity and
Community Stability.” clear impact on many others. Paine
AFTER developed the idea to include the
1994 In the US, a group of The keystone concept concept of “trophic cascades”—the
ecologists led by Brian Miller In the 1960s, Paine spent several strong, top-down effects that ripple
publishes a paper explaining years studying the animals of the through an ecosystem and its
the valuable role prairie dogs intertidal zone of Tatoosh Island organisms. Since Paine’s work
play as a keystone species. on the Pacific coast of Washington with starfish, several studies
State. He removed the ocher have demonstrated that there are
2016 Fieldwork leads marine starfish and watched its key prey,
ecologist Sarah Gravem to a mussel whose numbers had
conclude that organisms can Black-tailed prairie dogs look
been kept in check by the starfish, out from their burrow in a field in
be keystone species in some dominate the zone, replacing other Wyoming. Study of this species has
places but not in others. subordinate species. The removal revealed its key role in fostering
of a single, keystone species had a diversity in its native habitat.
See also: Predator–prey equations 44–49 ■ Mutualisms 56–59 ■ Animal ecology
106–113 ■ Trophic cascades 140–143 ■ Evolutionarily stable state 154–155
Whelks feed on They also display

barnacles; they aggression toward
are predators. their prey.
Robert Paine
In areas where
Whelks are like there are large Born in 1933, in Cambridge,
concentrations
little wolves in of barnacles, groups of
Massachusetts, Robert Paine
slow motion. studied at Harvard. After a
whelks congregate— stint in the US Army, where
like wolf packs. he was the battalion gardener,
Paine focused his research
on marine invertebrates.
His study of the relationship
many other keystone organisms, area to hunt for prey, and the ferrets between starfish and mussels
and they each fulfill their role in and tiger salamanders use the on the Paciic coast led him
to propose the concept
different ways. burrows for shelter. Almost 150
of keystone species—the
species of plant and animal are disproportionate impact that
Ecological engineers known to benefit from prairie dog a single species can have on
Prairie dogs in the American colonies. Although there are its ecosystem.
Midwest are a good example of a “losers”—notably vertebrates that Paine worked for most of
keystone species whose impact is favor tall vegetation—the prairie his career at the University
the result of their “engineering” dogs’ presence increases overall of Washington, where he
activities. Huge colonies of these biodiversity. When colonies die out, popularized field manipulation
small mammals dig networks of scrubby patches of mesquite experiments, or “kick-it-and-
tunnels beneath the prairie vegetation replace short grasses, see” ecology. He was awarded
grasslands. They sleep and raise plovers abandon the area, and the International Cosmos
their young in these extensive predator numbers decline. Award by the National
burrows, converting the grassland Academy of Sciences in 2013,
into a suitable habitat. Coral cleaners and died in 2016.
The prairie dogs’ constant The princess parrotfish in the Key works
digging dramatically increases Caribbean is another keystone
soil turnover and allows nutrients species, this time because of the 1966 “Food Web Complexity
and water from rain and snow to consequences of its feeding. The and Species Diversity,”
penetrate deeper than would fish lives around coral reefs, where American Naturalist
otherwise be the case. The damp, corals fight each other for light, 1969 “A Note on Trophic
nutrient-rich soil encourages a nutrients, and space. The parrotfish Complexity and Community
diversity of plants, and birds such scrapes the surfaces of the corals to Stability,” American Naturalist
as Mountain Plovers feed and nest remove layers of algal seaweed to 1994 Marine Rocky Shores
in the short grass. Predators like eat. If the parrotfish did not do this, and Community Ecology: An
Ferruginous Hawks and black- clumps of seaweed would grow on Experimentalist’s Perspective
footed ferrets are attracted to the the corals, smothering as well as ❯❯
64 KEYSTONE SPECIES
chemically damaging the reef. If the destructive behavior helps undigested seeds pass through
parrotfish was overfished or died maintain the feeding habitat for their gut, are then defecated, and
out from disease, the health of the grazing animals such as zebras, later germinate. Up to one-third
reefs would rapidly deteriorate. antelope, and wildebeest. It also of all West African tree species
indirectly helps the predators that depend on elephants for their seed
Landscape managers hunt the grazers—including lions, dispersal. Elephants also dig and
On African grasslands, elephants cheetahs, and hyenas—and the maintain waterholes, which benefit
smash down small and medium- smaller mammals that burrow in many other species.
sized trees for food, helping grassland soils. Without the Forest-dwelling Asian elephants
maintain savanna as grassland elephants, these animals would have a similar role. In southeast
and opening up new areas that soon disappear. Elephants are also Asia, they smash through gaps and
were formerly woodland. This very important seed dispersers; clearings in woodland, opening up
holes in the canopy. The new plants
that grow in these unshaded areas
Yellowstone wolfpack territories add to the forest’s plant and animal
diversity and also help a broader
range of animals to thrive there.
Cinnabar
Keystone predators
Junction
Butte The sea otter is a marine mammal
8 Mile
Prospect that lives in the Pacific coastal
Peak waters of North America. In the
Lamar 18th and 19th centuries, they were
Wapiti Canyon hunted extensively for their fur. By
Lake the early 20th century, they had
Cougar been wiped out in many areas, and
their total population was thought
Mollie’s to be fewer than 2,000 individuals.
Canyon
Since 1911, legal protection has led
to a slow increase in numbers.
Sea otters are important
because they eat large numbers
of sea urchins. These seafloor-
dwelling invertebrates graze on the
lower stems of kelp that grow up
Every species in the coastal

zone is influenced in one way
Bechler Snake River
or another by the ecological
effects of sea otters.
James Estes
American marine biologist
Each pack of wolves in the
Yellowstone National Park has its own
SCALE territory. Many of the territories overlap,
and numbers fluctuate from year to
10km (6 miles) year, with 108 wolves recorded in 2016.
Reintroducing Beavers were wiped out in the UK several dams on the headwaters
beavers to the UK 400 years ago, but the beneficial of the Tamar River, creating 13
role of this keystone mammal is new freshwater pools and making
now better understood. Beavers surrounding areas wetter.
are ecological engineers, building In Devon, the damp areas
dams and canals, and their created by beavers led to an
presence increases biodiversity. increase in the number of
In 2009, 11 beavers were bryophyte species (mosses and
reintroduced to Knapdale Forest, liverworts), and the range of
Scotland, and in 2011, the Devon aquatic invertebrates has risen
Wildlife Trust introduced a pair to from 14 to 41 species. Increased
a fenced enclosure. Both projects numbers of flying insects have
have been monitored to test their also improved bat diversity, with
impact on the environment. In two nationally rare bat species
Knapdale Forest, the beavers’ drawn into the area. More
dams changed the water level of beaver reintroduction projects
a loch, and Devon’s beavers built are now planned in the UK.
from the seabed, causing it to drift herbivores, such as beavers. Within fruited plant species share one or
away and die. If the kelp disappears, 10 years, the number of beaver two peaks of ripening each year.
however, so do the many other colonies increased from one to nine. Fig trees bear fruit throughout the
marine invertebrates that graze on Beaver dams helped revive wetlands, year, supporting many animals
it. “Forests” of kelp also absorb large and wetland wildlife flourished. The when other trees are fruitless.
amounts of atmospheric carbon increase in elk carcasses also More than 10 percent of the world’s
dioxide and, by slowing water benefited carrion-eaters—especially bird species and 6 percent of
currents, help protect coastlines coyotes, red foxes, grizzly bears, mammals (a total of 1,274 species)
from storm surges. The protection Golden Eagles, Ravens, and Black- are known to eat figs, as do a small
that sea otters offer kelp along billed Magpies—as well as several number of reptiles and even fish.
stretches of open coast is therefore smaller scavengers. Fig trees therefore provide a vital
particularly significant. Jaguars are apex predators in support mechanism for fruit-eating
Unlike the sea otter, some South and Central American forests, species. Without them, fruit bats,
keystone species are also “apex” preying on more than 85 species. birds, and other creatures would
predators at the top of the food Although there are very few jaguars decline or disappear. ■
chain, such as the gray wolf. Before in any given area, their impact on
1995, there had been no gray wolves the numbers of other predators—
in Yellowstone National Park for at such as caimans, snakes, large
least 70 years. American elk were fish, and large birds—as well as
common in the park, but there was herbivores, such as capybaras and
just a single colony of beavers. That deer, has a significant ripple-down
year, 31 wolves were introduced to effect on their ecosystem. Left By protecting a keystone
the park and by 2001 their numbers unchecked, the herbivores could species such as the prairie
had increased to more than 100, devour most of the plants and dog, the public could be
largely due to the abundance of destroy the habitat on which so educated about the value of
elk for food. many other species depend. ecosystem conservation.
The presence of wolves in the Brian Miller
park forced the elk to become more Keystone plants American ecologist
mobile. Rather than over-grazing Not all keystone species are
willow, aspen, and cottonwood trees animals. One example is the fig
in favored locations, the elk moved tree, of which there are about 750
on, allowing plants to regenerate species, mostly found in tropical
and provide a food source for other forests. In this habitat, most fleshy-
66
THE FITNESS OF
A FORAGING ANIMAL
DEPENDS ON ITS
EFFICIENCY
OPTIMAL FORAGING THEORY
E
very plant and animal helps predict the best strategy
IN CONTEXT on Earth needs resources that an animal can use to achieve
to survive. Plants obtain this goal.
KEY FIGURES
their nutrients and water from soil,
Ronald Pulliam (1945–),
and sunlight provides the energy Foraging theories
Graham Pyke (1948–), and
for photosynthesis. Animals The first theory of foraging by
Eric Charnov (1947–) generally have to work harder to wild animals did not emerge until
BEFORE find their food—they have to move, the mid-1960s, when Americans
1966 John Merritt Emlen, and this uses extra resources. Robert MacArthur and Eric Pianka
Robert MacArthur, and Eric Optimal foraging theory (OFT) examined the question of why,
Pianka outline the concept proposes that animals try to gather when a range of food was available
of optimal foraging in two resources in the most efficient way to them, animals often restricted
articles published in the to avoid using additional energy. themselves to a few preferred types
Searching for and capturing food of prey. They argued that natural
American Naturalist magazine.
takes energy and time. The animal selection favored animals whose
AFTER needs to gain maximum benefit behavior maximized their net
1984 Argentinian–British for minimal effort in order to energy intake per unit of time spent
zoologist Alejandro Kacelnik achieve optimal fitness. OFT foraging. An animal’s foraging time
researches the foraging includes searching for prey and the
behavior of starlings to killing and eating of the food
illustrate the marginal (handling time).
value theorem (MVT). These ideas were developed by
American ecologists Ronald Pulliam
1986 Belgian ecologist Patrick and Eric Charnov and Australian
Meire investigates prey Diets should be broad ecologist Graham Pyke. It seems
selection by oystercatchers. when prey are scarce, that OFT works best for mobile
but narrow if food foragers seeking immobile prey, and
1989 Swiss environmental is abundant. some researchers believe it is less
scientists T. J. Wolfe and Paul Eric Pianka relevant when prey are mobile.
Schmid-Hempel examine how
the weight of nectar carried Key choices
by bees has an effect on the Animals must choose which types
bees’ foraging behavior. of food to eat, which is rarely
straightforward. For example,
See also: Evolution by natural selection 24–31 ■ Predator–prey equations
44–49 ■ Competitive exclusion principle 52–53 ■ Mutualisms 56–59
small clams was better spent

digging for another, larger clam.
A similar study with oystercatchers
and mussels found that the largest
mussels were left—they had
The expected behavior of thicker, barnacle-clad shells, so
animals with respect to opening them was more difficult.
available resources can be The oystercatchers benefited more
used to predict … the biotic by looking for thin-shelled mussels,
structure … of communities. despite their smaller size.
Ronald Pulliam Animals also have to make Echolocating bats
choices about where and when to
feed. The longer a starling spends Technological advances have
in one patch of suitable grassland, greatly helped research into
for example, the harder it will the hunting strategies of
become to find prey, so it has to animals. Insectivorous bats
decide when to abandon that patch (also known as microbats)
American ecologists Howard and move to another—an example use echolocation in the
Richardson and Nicolaas Verbeek of what is known as the “marginal dark to locate and pursue
studied Northwestern Crows value theorem.” Foraging animals flying insect prey, such as
feeding on clams in the intertidal also need to consider a range of moths and midges. A team
zone of British Columbia. The other factors such as the presence of Japanese scientists set out
to study the bats’ feeding
crows put lots of effort into digging of predators, the number of animals
behavior using microphone
clams out of the mud, opening the competing for the same food, and
array measurements and
shells, and feeding on the animal the impact of human activity. ■ mathematical modeling
inside. The ecologists noticed that analysis. The researchers
smaller clams went unopened and recorded the echolocation
Oystercatchers, despite their name,
concluded that the crows had to are reliant on cockles and mussels as calls and flight paths of the
make an energy trade-off between their primary food source. Without bats and discovered that they
handling time and edible food. The these shellfish, they are forced to often directed their sonar not
time and energy needed to open up forage farther inland. just at their immediate prey
but at the next target they
were lining up as well.
The team also found
evidence that the bats chose
flight paths that would allow
them to plan two steps ahead,
rather like skilled chess
players. Not only were the
animals maximizing their
energy input by targeting
multiple prey items, but they
were also minimizing their
energy output by reducing
the distance they flew in
pursuit of insects. This
behavior fits in well with
optimal foraging theory.
68
IN CONTEXT
PARASITES AND KEY FIGURES

Roy Anderson (1947–),
PATHOGENS CONTROL
Robert May (1936–)
BEFORE
1662 English statistician John
POPULATIONS LIKE
Graunt seeks to classify causes
of death in London in Natural
PREDATORS
and Political Observations
made upon the Bills of Mortality.
1927 Scottish scientists
ECOLOGICAL EPIDEMIOLOGY
Anderson Gray McKendrick
and William Ogilvy Kermack
develop an epidemic model for
infected, uninfected, and
immune individuals.
AFTER
1996 American epidemiologist
James S. Koopman calls for
greater use of computational
technologies to simulate
disease generation and spread.
2018 A global team tracks the
origins and spread of a fungus
devastating frog populations.
E
pidemiology is the study
of how disease spreads
through a population. Its
initial application was to human
diseases, but its methods have
been recognized as an effective
way of modeling populations of
other organisms, too.
Ecologists have long known
that the size of an animal or plant
population and its growth rate
depend on the availability of food,
living space, and levels of
predation. In the 1970s, British
epidemiologist Roy Anderson and
Australian scientist Robert May
showed how parasites and
infections from pathogens such as
bacteria and viruses limited the
See also: The microbiological environment 84–85 ■ Microbiology 102–103 ■ The ubiquity of mycorrhizae 104–105
■ Biodiversity and ecosystem functioning 156–157
Map of deaths from cholera in London in 1854

KEY
1–4 deaths Street
Oxford
5–9 deaths Soho

Square
10–15 deaths
Broad Street
pump
eet
d Str
B r oa
Re
ge
nt
S
t re
Fatalities in London’s et
et
cholera outbreak of Golden re

St
1854 were linked to the Square er
central pump; its water ew
Br
was found to have
been contaminated
with infected sewage
from a stricken family.
size of a population. In wild sheep, thought to be caused by miasma—

for instance, the chief cause of a sort of poisonous vapor in the
death is lungworms, while most air—that spread from the bodies of
wild birds die from viral infections. the dead and dying. Snow was not
In ecology, the effects of disease the first to question this theory, but
have wider implications. Up to he was especially suspicious of it in
40 percent of ocean bacteria are the case of cholera.
killed each day by viruses. This In 1854, Snow plotted every
causes a “viral shunt,” because case of cholera on a map of Soho,
nutrients that would otherwise flow and found that afflicted households
up the food chain to consumers collected their water from a pump
revert to the bottom of the chain. on Broad Street (later renamed
Broadwick). He shut down the ❯❯
Human beginnings
Epidemiology has its beginnings
British doctor John Snow fought the
in the work of physician John Snow, establishment to gain acceptance for
who witnessed a cholera epidemic his belief that cholera was waterborne.
in the Soho district of London in The medical journal The Lancet finally
1854. At the time, disease was conceded that he was right in 1866.
70 ECOLOGICAL EPIDEMIOLOGY
The role of drought pump, and the epidemic soon
in plant diseases ended. This showed that cholera
was a waterborne disease that
Like other disease-causing humans contracted through
agents, a plant pathogen contaminated food and drink. A
(disease-causing agent)
decade later, Louis Pasteur’s “germ
needs a supply of susceptible
individuals to infect. Periods theory” proposed that diseases, as
of drought slow the rate of well as general rotting and decay,
plant reproduction and were the work of microorganisms.
growth, thereby reducing the
prevalence of disease. Disease model
Aridity, however, also In their 1970s studies, Anderson
weakens plants and makes and May focused first on building
them susceptible to pathogens a mathematical model to show
that thrive in dry conditions. how a microorganism could affect
These include various forms a population. This led to a set of
of fungi that attack the leaves equations that they hoped would
of grain crops, legumes, and help explain the real-life impact of A ravaged tree in North Yorkshire,
fruits. These fungi are adapted different kinds of pathogens, from UK, shows the effects of Dutch elm
to survive in a dormant state disease, a fungus spread by elm bark
bacteria and viruses to parasitic
as hardened microscopic beetles accidentally introduced to
worms and insect larvae.
bodies in soil. They can exist Europe and America from Asia.
for many years in dry soil, but In their model, a number of mice
when the soil becomes wet, were divided into three groups:
the fungi must find a host susceptible (uninfected) mice, of disease, the total would remain
within a few weeks or die. infected mice, and mice that had more or less the same, with the
They do not necessarily kill survived infection and were now rate of added mice balancing that
their host. Recent research immune. Unlike many earlier at which other mice died.
into chickpeas suggests that epidemiological models, the total For simplicity, the model
although infections from such population was not a fixed number; assumed that the diseases were
fungi do increase during a dry mice could be added either by transmitted by contact between
spell, the mortality rate of the reproduction or by additions from infected and uninfected mice. Not
affected plants goes down other populations. Mice also died all infected mice would die, so the
during a drought. from natural causes. In the absence model also included a recovery rate.
Mice that recovered would be
immune, at least initially. Immunity
to viruses is more or less lifelong,
but it is possible to become
susceptible again to the same
bacterial infection as time passes.
Sensibly used, Therefore, the calculations also
mathematical models are included a rate of loss of immunity.
no more and no less than Putting all this together,
tools for thinking about Anderson and May produced a
things in a precise way. set of equations to predict the rate
Roy Anderson and of population change in the three
Robert May initial groups of uninfected but
A summer drought produces susceptible mice, infected mice,
only sparse growth of young barley and the immune survivors. These
plants. Lack of moisture and too
much heat reduce their resistance equations could be added together
to fungi that attack their roots. to give the rate of change for the
total mouse population.
effects of hypothetical values. They by the disease. Infection numbers
found, for instance, that when the rise sharply to a maximum, then
rate of added mice was highest, the drop away. Epidemics also occur
disease had the greatest impact on when a disease is not particularly
population numbers. This suggests deadly but slows the population
Diseases such as that species with high reproductive growth rate; this has occurred with
measles and rubella, rates (introducing large numbers human diseases such as measles
with short infections of uninfected offspring) are most and chickenpox.
and lasting immunity, likely to have endemic diseases
will tend to exhibit within the population, and show Applying the theory
epidemic patterns. depressed numbers compared with The characteristics of disease
Roy Anderson species that breed more slowly. and its effects on animal and
They also explored the differing plant populations are of increasing
effects on populations of diseases ecological importance. Food
of different intensities. producers, for example, benefit from
Unlike endemic diseases, studies into the nature of parasites
in which the population’s level and the dynamics of diseases that
of infection remains consistent, can affect crops and livestock.
From their calculations, they epidemics appear in populations Conservationists also employ
deduced that a disease will persist when the growth rate of all infected epidemiology to predict how exotic
in a population whose equilibrium and uninfected members is low diseases and invasive parasites
point (the rate of new additions, compared to the death rate caused might affect fragile ecosystems. ■
balanced by the natural death rate)
is greater than the combined
effects of natural mortality, disease Venn diagram of ecological epidemiology
deaths, recovery, and transmission
Susceptible host
rate. While the disease is present,
that equilibrium point will be lower
than if the population were disease
free. If, however, the equilibrium
point of a population affected by
no
disease is lower than the combined disease
effects of deaths, recoveries, and
rate of transmission, the disease
will die out. Once a population is no no
disease free, its equilibrium point disease disease
will return to its former level.
disease
Matching the real world
Anderson and May needed to show
that their model was an accurate no no no
predictor of a real-life population. disease disease disease
They did so by using data from
a study of laboratory mice infected
with the bacterial disease Pathogen Favorable
pasteurellosis; the data included environment
the impact on the population of for pathogen
adding individuals at different A pathogen strikes when it finds a suitable host in an
rates. The observed data confirmed environment that favous infection, as shown where the
their predictions, so the two circles intersect. For instance, diarrheal diseases spread
scientists were able to consider the quickly among sick people in unsanitary conditions.
72
WHY DON’T
PENGUINS’
FEET FREEZE?
ECOPHYSIOLOGY
T
he central principle of to its distribution, abundance, and
IN CONTEXT Darwinian evolution is fertility. Ecophysiology now plays
that all organisms, from an important role in helping
KEY FIGURE
simple bacteria to complex scientists understand how the
Knut Schmidt-Nielsen
mammals, are adapted by natural stresses created by climate change
(1915–2007)
selection to survive in a particular impact on both wild ecosystems
BEFORE niche and habitat. Ecophysiology— and cultivated environments.
1845 The explorer Alexander for which Knut Schmidt-Nielsen’s
von Humboldt reveals that book Animal Physiology (1960) was Managing temperature
plants facing similar ecological a vital inspiration—is the study of Ecophysiology has revealed a
factors also have many an organism’s anatomy and how it number of specific adaptations
analogous features. functions (its physiology), as well as for different environments. For
how these characteristics relate to example, animals that live in colder
1859 Charles Darwin argues the challenges posed by its regions tend to have larger bodies
that organisms evolve because environment. It shows how the and smaller legs, ears, and tails
they are adapting to changed anatomy of an animal or plant is than related species living in
ecological conditions. linked to its ability to survive, and warmer climes. A larger body has
a smaller surface-area-to-mass
AFTER ratio, and therefore loses heat more
1966 Australian biochemists slowly, while smaller appendages
Marshall Hatch and Charles reduce exposure to frostbite.
Slack explain that the most In the most extreme cold, the
widespread plants are the feet of a warm-blooded animal
ones that photosynthesize From a physiological are at risk of becoming frozen to
most efficiently. viewpoint, freshwater the ground. Mammals in Arctic
is no more freely regions such as musk oxen and
1984 Peter Wheeler, a British available in the sea polar bears are adapted for life in
scientist, suggests that human than in the desert. these conditions by having thick
bipedalism—the ability to Knut Schmidt-Nielsen hairs to insulate their feet.
walk on two legs—evolved as In the Antarctic, the undersides
a thermoregulatory adaptation of penguins’ feet are insulated
that reduces the body’s by a thick layer of fat. Penguins
exposure to direct sunlight. also have a heat-exchange (or
counter-current) mechanism in
See also: Evolution by natural selection 24–31 ■ Ecological niches 50–51
■ Competitive exclusion principle 52–53 ■ Ecological stoichiometry 74–75
their legs. The warm blood arriving cooler than the air outside, so the
from the body is cooled to near moisture it carries condenses in
32°F (0°C) by the chilled blood the nose. This creates the cool,
arriving from the feet, which damp conditions needed to chill
warms to body temperature in the next in-breath.
the process.
Gazelles in Africa use a similar Future challenges
counter-current system to cool their Today ecophysiology is becoming
body temperature. They are able to increasingly focused on plants,
chill the blood entering their head, fungi, and microbes. Like animals,
giving them an advantage over they have to adapt to survive—and Knut Schmidt-Nielsen
their predators, who often overheat. studying them offers the possibility
Camels have a heat-exchange of vital discoveries for commercial Knut Schmidt-Nielsen grew
system in their nasal cavity, which and conservation purposes. ■ up in the Norwegian town of
reduces the amount of water lost in Trondheim. His interest in the
their breath. Hot, dry air is inhaled Emperor penguins survive freezing way animal physiology related
and mixes with moisture inside Antarctic temperatures thanks in part to habitat was inherited from
the nose before traveling to the to the way their bodies have evolved his grandfather who, years
lungs. The exhaled air is much to adapt to the harsh environment. before Knut’s birth, had
released thousands of flounder
(a marine fish) hatchlings into
a freshwater lake. Although
the fish thrived, they were
unable to breed because their
reproductive physiology was
adapted for life in salt water.
Schmidt-Nielsen joined
Duke University, North
Carolina, in 1954. He built a
climate-controlled space for
keeping desert animals, where
he considered the anatomy
of camels, gerbils, and other
species able to live for long
periods without water. He also
investigated the respiratory
systems of birds and the
buoyancy of fish. His 1960
textbook Animal Physiology
is still a classic work.
Key works
1960 Animal Physiology

1964 Desert Animals
1972 How Animals Work
1984 Scaling
1998 The Camel’s Nose:
Memoirs of a Curious Scientist
74
ALL LIFE IS
CHEMICAL
ECOLOGICAL STOICHIOMETRY
E
very living organism— The field of ecological stoichiometry
IN CONTEXT from tiny ocean algae was comprehensively described
to a mighty redwood—is for the first time by American
KEY FIGURES
made up of chemical elements biologists Robert Sterner and James
Robert Sterner (1958–),
in varying ratios. Ecological Elser; in Ecological Stoichiometry
James Elser (1959–)
stoichiometry considers the (2002), they used mathematical
BEFORE balance of these elements, and models to demonstrate the
1840 German biologist and how the ratios change during application at every level, from
chemist Justus von Liebig chemical reactions. Studying molecules and cells to individual
asserts that the limitations such ratios throws light on the plants and animals, populations,
on agriculture productivity are way the living world operates, communities, and ecosystems.
primarily chemical. revealing how organisms obtain
the nutrients and other chemicals Key chemicals
1934 US oceanographer Alfred they require for life from the In ecological research, the three
Redfield measures the atomic resources in their environment. main elements examined are
ratio of carbon, nitrogen, and carbon (C), nitrogen (N), and
phosphorus (C:N:P) in plankton phosphorus (P), because each
and seawater, and finds it to be plays a vital role. Carbon is a basic
relatively consistent in all building block of all life and an
oceans. The Redfield Ratio soon important part of many chemical
becomes a benchmark for such processes. Nitrogen is a major
research in all habitats. Individual organisms also constituent of all proteins, while
show differences in phosphorus is crucial for cell
AFTER stoichiometry during their life development and storing energy.
2015 In “Ocean stoichiometry, cycles. Young organisms may An organism’s C:N:P ratio is not
global carbon, and climate,” have different compositions necessarily consistent. Plants have
Robert Sterner highlights from older ones … a variable ratio: they can adjust the
inconsistencies in C:N:P ratios Robert Sterner and balance of their elements according
in phytoplankton, which absorb James J. Elser to their environment. For instance,
more atmospheric carbon in the proportion of carbon in their
low-nutrient, low-latitude ocean chemical makeup can rise on a
surface waters and adjust their particularly sunny day because
ratios accordingly. more photosynthesis occurs—the
process by which they take carbon
See also: Ecophysiology 72–73 ■ The food chain 132–133 ■ Energy flow through
ecosystems 138–139 ■ The foundations of plant ecology 167
The Growth
Rate Hypothesis
Cancer research is one area
Controlling ecological stoichiometry ratios where stoichiometry is now
A locust eats grass that may contain six times being employed. Evidence is
as much carbon as it needs. To get the right growing for a theory called the
balance, it excretes carbon or breathes it out Growth Rate Hypothesis
as CO2. Locusts are widely used in research (GRH), which may help explain
because they are easy to breed. why some cancerous tumors
grow at faster rates than the
rest of the body.
The hypothesis states that
organisms with high C:P
KEY (carbon:phosphorus) ratios,
LOCUSTS GRASS such as fruit flies, have more
Carbon Nitrogen 5:1 33:1
ribosomes in their cells, which
enables them to grow and
dioxide from the air and use the may adjust its digestive enzymes reproduce more rapidly.
sun’s energy to convert it into the and excrete it, store it as fats, or raise Around half of all phosphorus
in an organism is in the form
nutrients they require. its metabolic rate to burn it off,
of ribosomal RNA (rRNA); it is
Higher up the food chain, breathing out the excess carbon as
present in every cell, creating
animals have largely fixed C:N:P CO2. Overuse of such mechanisms proteins to build new cells and
ratios, so they must deploy various to redress a high imbalance can, grow the body. Applying
mechanisms to deal with any however, affect fitness, growth, and biological stoichiometry,
imbalances of chemicals entering reproduction. An animal that eats James Elser and his team
the body. If an insect or animal other animals has less work to do, have shown that fast-growing
herbivore is getting too much carbon because its prey’s C:N:P ratio tumors have a much higher
from its plant diet, for instance, it closely matches its own. However, phosphorus content than
the size of its prey population is still normal body tissue. Such
determined by the plants in its research may help scientists
environment because plants with a understand how tumor growth
high carbon ratio can only support could be controlled.
a small food chain of consumers.
Understanding our world

Food chains are one area of study;
ecological stoichiometry covers just
about everything and all the links
in between. By discovering how
the chemical content of organisms
shapes their ecology, scientists are
also learning how environments
can be better managed. Their
findings may significantly influence
the future of life on Earth. ■
Malignant lung tissue (seen

The desert locust (Schistocerca here) and cancerous colon tissue
gregaria) has to eat vast quantities both had the highest phosphorus
of carbon-rich plants in order to get content in research exploring the
enough nitrogen and phosphorus rapid growth rates of tumors.
to maintain its C:N ratio.
76
FEAR ITSELF
IS POWERFUL
NONCONSUMPTIVE EFFECTS OF
PREDATORS ON THEIR PREY
M
any descriptions of risks of being eaten. The lethal role
IN CONTEXT ecosystems focus on of predators is obvious, but their
predator–prey interactions nonlethal (nonconsumptive) role
KEY FIGURE
in which predators kill and prey are can have an even bigger impact on
Earl Werner (1944–)
eaten. However, American ecologist an ecosystem. Potential prey are
BEFORE Earl Werner and others have shown forced to change their way of life
1966 American ecologist that the mere presence of a predator in order to avoid being killed.
Robert Paine conducts a affects the behavior of prey. In 1990, Werner studied the
series of groundbreaking Apart from apex predators, all effects of green darner dragonfly
field experiments to highlight animals must balance the need to larvae on toad tadpoles. He noticed
the crucial effects of a predator sleep, reproduce, and feed with the that when the predatory larvae
on the community in which
it lives.
1990 Canadian biologists
Steven Lima and Lawrence In the presence of predators …
Dill analyzed the decision-
making of organisms that are
at the greatest risk of being
preyed on by other creatures.
AFTER prey move on to prey spend more time
2008 American behavioral other areas even if there is hiding in sheltered habitats
biologist and ecologist John less food there. than feeding in the open.
Orrock teams up with Earl
Werner and others to produce
mathematical models to
explain the nonconsumptive
effects of predatory animals. Even without preying on
them, predators can cause prey to
fail to thrive.
See also: Evolution by natural selection 24–31 ■ Predator–prey equations 44–49 ■ Ecological niches 50–51
■ Competitive exclusion principle 52–53 ■ Mutualisms 56–59 ■ Optimal foraging theory 66–67
bullfrog’s new behavior gave it

a competitive advantage over the
green frog by making it bigger.
Terrestrial animals
Early studies of nonconsumptive
effects (NCEs) were concerned with
aquatic organisms under laboratory
conditions, but more work has now
been done in the wild with land-
dwelling animals. German field
research published in 2018
focused on lynx and their roe deer
prey. When lynx were present,
researchers found that the roe deer
avoided areas they knew to be
high-risk, both during the day and
on summer nights when nocturnal
predation is more common. The
deer treated some grazing areas as
out of bounds, presumably due to
fear of being attacked by lynx.
Wherever there are predators,
they exert NCEs. They also affect
some sessile (nonmoving) species,
as well as mobile prey. This can
happen when certain dominant
competitors are displaced by
predators and, in their new
A green darner dragonfly laying when predatory dragonfly larvae habitats, outcompete sessile
its eggs in a pond. The larvae that were introduced to the tank, both animals for food. Small fish that
hatch out are predators and have been prey species became less active are displaced, for example, could
shown to influence the behavior of their
tadpole prey.
and chose different places in which outcompete sponges for food. ■
to swim. The bullfrog tadpoles grew
more quickly than they had in a
were in the tank, the tadpoles were predator-free tank, but the green
less active, swam to other parts of frog tadpoles decreased their
the tank, and metamorphosed into feeding activity and grew more
adults when they were smaller. The slowly. Werner concluded that for
predator had changed the toads’ prey species there was a trade-off … species react [to predators]
morphology and their behavior just between the need to grow as fast by reducing activity and
by being there. as possible and the risk of predation. altering space use.
In 1991, Werner investigated Growing more quickly requires Earl Werner
what happened when more than more feeding activity, and this in
one prey species was involved. In turn increases the chances of being
the absence of a predator, bullfrog eaten by a predator. As the larvae’s
and green frog tadpoles grew at presence altered the behavior of
virtually identical rates. However, the prey species differently, the
ORDERIN
NATURAL
G THE
WORLD
80 INTRODUCTION
Aristotle’s History of A private collection of natural The Natural History

Animals groups living things history curiosities is displayed Museum in London opens its
based on their species, in a at Oxford University’s doors to the public, free of
scala naturae that places Ashmolean Museum, the charge. It now houses 80
organisms into 11 grades. world’s first public museum. million specimens.
C.350 BCE 1683 1881
1665 CE 1758
Micrographia, the richly The 10th edition of Systema

illustrated book by Robert Naturae by Carl Linnaeus
Hooke, reveals microscopic classifies a range of plant
structures to a wider audience. and animal species using
his binomial system.
P
eople have long marveled at In keeping with the prevailing to adopt this new technology: his
the variety of life, celebrating ideas of the Catholic Church, the book Micrographia (1665) inspired
nature’s gifts in prehistoric natural world was seen as static others to do likewise. Able to view
cave art that dates back 30,000 years and unchanging. specimens magnified to 50 times
or more. In Ancient Greece in the 4th their actual size, he made meticulous
century BCE, Aristotle made an early An age of discovery drawings of microsopic life, and
attempt to classify living organisms; The age of great expeditions also coined the term “cell” after
his 11-grade scala naturae (“ladder of of discovery revealed previously examining plant fibers. Hooke also
life”) placed humans and mammals uncharted regions and their suggested a living origin for fossil
at the top, and descended through animals and plants. In his History fragments found in rocks.
other, more “primitive” animals of the Animals (1551–58), Swiss
to plants and then minerals. A physician and naturalist Conrad Classifying variety
thousand years later, the medieval Gesner included some of the recent English vicar John Ray’s History of
world still considered variations finds from the New World and the Plants (1686–1704) was the botanical
on Aristotle’s system to be valid. Far East, as well as relying on equivalent of Gesner’s earlier work,
There were several reasons for this. classical literature. The five-volume listing some 18,000 species in three
Without microscopes, nothing was work reflected his division of huge volumes. Ray also produced
known of cells and microorganisms. animals into mammals; reptiles and a biological definition of a species,
Without the means to explore amphibians; birds; fish and aquatic remarking that “one species never
underwater, science’s knowledge animals; and snakes and scorpions. springs from the seed of another.”
of aquatic creatures was limited, The invention of the microscope Swedish botanist Carl Linnaeus,
and many parts of the world were also had a major impact. English the “father of taxonomy,” first
still unknown to Western scientists. scholar Robert Hooke was quick published Systema Naturae in 1735,
ORDERING THE NATURAL WORLD 81
Norman Myers’s
Carl Woese “biodiversity hotspots”
establishes a new, third concept identifies ten hotspots
category of organisms— where conservation efforts
the prokaryotes. should preserve rare species.
1977 1988
1942 1988 2018
Ernst Mayr develops the Edward O. Wilson coins The IUCN Red List shows that
biological species concept, the term biodiversity more than 26,000 species –
which categorizes species and later identifies the more than 27 percent of all
based on their ability to key human threats those assessed—are at
breed with each other. to biodiversity. risk of extinction.
but it is the 10th edition from 1758 but a population that can breed new domains. As of 2018, about
that founded the modern zoological only among themselves. Mayr went 1.74 million extant plant and animal
naming system. Two volumes of on to explain how if groups within species have been described, but
Linnaeus’ work are devoted to a species become isolated from the estimates of the total number range
plants and animals, which he rest of the population, they may from 2 million to 1 trillion.
divided into classes, orders, genera, start to differ from the rest, and
and species. The binomial system, over time, through genetic drift and The threat to diversity
in which every species is given a natural selection, may even evolve By the late 20th century, however,
generic name followed by a specific into new species. alongside a growing knowledge
name, is still in use today. Linnaeus Modern technological advances, of the scale and critical role of
also wrote a third volume on rocks, including electron microscopy and biodiversity—and of how evolution
minerals, and fossils. mitochondrial DNA analysis, have can destroy species as well as
revealed much information—some create them—American ecologist
Species concepts of it surprising—about the number E.O. Wilson and others made the
Building on Darwin’s theory of of species and the relationships world aware that human activity
evolution by means of natural between them. In 1966, striving to was responsible for causing a rapid
selection, German-American reflect the intricacies of evolution, acceleration in the extinction rate.
evolutionary biologist Ernst Mayr German entomologist Willi Hennig Some have even warned that Earth
cemented the biological concept of proposed a new taxonomic system could be on the verge of a sixth
species in his Systematics and the of clades—groups of organisms mass extinction. Many policies are
Origin of Species (1942). He argued based on a common ancestor. In now being proposed to counter
that a species is not just a group of the 1970s, American biologist Carl this, including the protection
morphologically similar individuals, Woese classified all life into three of biodiversity hotspots. ■
82
IN ALL THINGS
OF NATURE THERE
IS SOMETHING
OF THE MARVELOUS
CLASSIFICATION OF LIVING THINGS
F
rom the beginning of Aristotle placed animals in a scala
IN CONTEXT recorded history, people naturae (ladder of nature), with 11
have attempted to identify grades distinguished by their mode
KEY FIGURE
organisms according to their uses. of birth. Those in the top grades
Aristotle (c. 384–322 bce)
Egyptian wall paintings from gave birth to live, hot, wet offspring;
BEFORE c. 1500 bce show, for example, that those in the lower grades to cold,
c. 1500 bce Different people understood the medicinal dry eggs. Humans were at the very
properties of plants are properties of many plants. In the top of the scale, with live-bearing
recognized by ancient text History of Animals, written tetrapods (four-legged creatures),
Egyptians. in the 4th century bce, the Greek cetaceans, birds, and egg-laying
philosopher and scholar Aristotle tetrapods lower down. Aristotle
AFTER made the first serious attempt to placed minerals on the bottom
8th–9th centuries ce Islamic classify organisms, studying their grade of his scale, with plants,
scholars of the Umayyad and anatomy, life cycles, and behavior. worms, sponges, larva-bearing
Abbasid dynasties translate insects, and hard-shelled animals
many of Aristotle’s works Features of classification on the levels above.
into Arabic. Aristotle divided living things into
plants and animals. He further
1551–58 Conrad Gessner’s grouped about 500 species of
History of Animals classifies animals according to obvious
the animals of the world into anatomical features, such as
five basic groups. whether they had blood, were
“warm-blooded” or “cold-blooded,” If any person thinks the
1682 John Ray publishes his examination of the rest
whether they had four legs or more,
History of Plants, which lists of the animal kingdom
and whether they gave birth to live
more than 18,000 species. an unworthy task, he must
offspring or laid eggs. He also noted
1735 Carl Linnaeus devises a whether animals lived in the sea, hold in like disesteem
system of binomial names, the on land, or flew in the air. Most the study of man.
first consistent classification of significantly, Aristotle used names Aristotle
organisms, according to which for his groupings that were later
he names every species listed translated into the Latin words
in his Systema Naturae. “genus” and “species”—terms
that are still used by modern
taxonomists to this day.

See also: The microbiological environment 84–85 ■ A system for identifying all nature’s organisms 86–87 ■ Biological
species concept 88–89 ■ Microbiology 102–103 ■ Animal behavior 116–117 ■ Island biogeography 144–149
An octopus blends in with its

surroundings. The ability of these
creatures to change color was one of
Aristotle’s many accurate observations.
While Aristotle’s system of

classification was rudimentary, it
was based largely on first-hand
observations, many of which were
made on the island of Lesvos. He
recorded things that noone else
had described, including that
young dogfish grew inside their
mothers’ bodies, male river catfish
guard eggs, and octopuses
can change color. Most of his
observations were good—and some
were confirmed only centuries later.
The great chain of being The Swiss doctor Conrad Gessner (reptiles and amphibians); birds;
Despite its limitations, Aristotle’s wrote the first modern register of fish and aquatic animals; and
method of classification heavily animals—also called History of snakes and scorpions. In 1682,
influenced every later attempt at Animals—in the mid-16th century. the English naturalist John Ray
grouping animals and plants until This monumental five-volume work produced the equivalent register
the 18th century. Medieval was based on classical sources but for botany with his History of
Christianity developed his scala included newly discovered species Plants. Within little more than
naturae as a “great chain of being,” from East Asia. It covered the main 50 years, the classification of
with God at the top of a strict animal groups as Gessner saw living things would be completely
hierarchy, humans and animals them: live-bearing quadrupeds transformed by Carl Linnaeus’s
beneath, and plants at the bottom. (mammals); egg-laying quadrupeds Systema Naturae. ■
Aristotle Aristotle was born in Macedonia, scholar Ptolemy and King

ancient Greece. Both his parents Alexander the Great. In 335 bce,
died when he was young, and he he established his own school
was raised by a guardian. Aged at the Lyceum in Athens. After
17 or 18, Aristotle joined Plato’s Alexander’s death in 322 bce,
Academy in Athens, where he Aristotle fled the city, and died
studied for 20 years, writing on on the island of Euboea in the
physics, biology, zoology, politics, same year.
economics, government, poetry,
and music. Later, he traveled Key works
to the island of Lesvos with a
student named Theophrastus 4th century bce
to study the island’s botany and History of Animals
zoology. Much of his History On the Parts of Animals
of Animals was based on On the Generation of Animals
observations he made there. On the Movement of Animals
Aristotle taught both the future On the Progression of Animals
84
BY THE HELP
OF MICROSCOPES
NOTHING ESCAPES
OUR INQUIRY
THE MICROBIOLOGICAL ENVIRONMENT
L
eafing through the pages of Although it is not known for
IN CONTEXT Micrographia, a 17th-century certain who developed the first
reader would have been microscopes, they were certainly
KEY FIGURE
astonished. Here, in English in use by the 1660s. The early
Robert Hooke (1635–1703)
scientist Robert Hooke’s seminal instruments were unreliable—due
BEFORE 1665 book, were many detailed to the difficulty of making the
1267 English philosopher illustrations of structures previously lenses—and scientists had to be
Roger Bacon discusses the use hidden from the human eye due to inventive and work around the
of optics for looking at “the their minuscule size. Hooke’s problem. At first, Hooke had
smallest particles of dust” in microscope magnified things by a difficulty seeing his specimens
his Opus Majus Volume V. factor of fifty, but the accuracy of clearly, so he invented an improved
his drawings also owes much to light source, named a “scotoscope.”
1661 Microscopic drawings by his painstaking approach. Hooke Hooke’s book is more than just
English architect Christopher would make numerous sketches an accurate representation of what
Wren impress Charles II, who from many different angles before he saw through the lens; it also
commissions more drawings combining them into a single image. theorizes on what the images reveal
from Robert Hooke. about the workings of the organisms
he studied. For example, when
AFTER looking at a wafer-thin specimen of
1683 Dutch amateur scientist cork, Hooke saw a honeycomb-like
Antonie van Leeuwenhoek pattern, the elements of which he
uses a microscope to observe described as “cells”—a term that is
bacteria and protozoa, and … in every little particle… still used today.
publishes his findings with we now behold almost as
the Royal Society of London. great a variety of Creatures, Microscopic marvels
as we were able before to Micrographia inspired many
1798 Edward Jenner, an reckon up in the whole other scientists to investigate the
English physician and Universe itself. microscopic world. Following
scientist, develops the world’s Robert Hooke notes and diagrams from Hooke’s
first vaccine—for smallpox— book, Dutch scientist Antonie
and publishes An Inquiry into van Leeuwenhoek was able to
the Causes and Effects of the construct his own microscopes.
Variolae Vaccinae. He achieved magnifications of
more than 200 times actual size.

See also: Classification of living things 82–83 ■ A system for identifying all nature’s organisms 86–87 ■ Microbiology
102–103 ■ Thermoregulation in insects 126–127
Van Leeuwenhoek examined

samples of rainwater and stagnant
pondwater and marveled at the
multitude of life he saw there. He
identified single-celled protozoa,
naming them “animalcules,” and [Micrographia is] … the
went on to discover bacteria. He most ingenious book that I
also made many observations of ever read in my life.
human and animal anatomy, Samuel Pepys
including blood cells and sperm. English diarist
While van Leeuwenhoek
examined water samples, fellow
Dutchman Jan Swammerdam was
placing insects under his own
microscope. He published records
of all manner of insects depicted in
the finest detail and uncovered organ. Grew also spotted pollen
much about their anatomy. grains and noted that they were
Swammerdam’s most influential transported by bees.
work was Life of the Ephemera Since the early days of
(1675), which recorded in great microscopy, devices have grown
detail the life cycle of the mayfly. in sophistication. The electron
In England, Nehemiah Grew microscope, first used in 1931, uses
used microscopy to examine a beams of electrons—rather than
wide range of plants. He was the light—to reveal objects, allowing
first to identify flowers as being scientists an even closer look. The compound eye and brain of a
bee, drawn by Jan Swammerdam and
the sexual organs of plants. In The Electron microscopes provide views published in A Treatise on the History
Anatomy of Plants (1682), Grew of up to one million times actual of Bees, shows the eye exterior (left)
named the stamen as the male size—600 times greater than most and the eye dissected (right), with the
organ and the pistil as the female modern light microscopes. ■ brain cross-sectioned below.
Robert Hooke Born on the Isle of Wight, England, include some early insights into
Hooke showed an early interest the wave theory of light; the
in science. A small inheritance construction of some of the
allowed him to attend the earliest telescopes; and the
prestigious Westminster School, formulation of Hooke’s Law.
where he excelled, earning a place Hooke was also a respected
at Oxford University. There he architect, an activity that made
assisted the natural philosophers him a wealthy man.
John Wilkins and Robert Boyle.
In 1662 Hooke became the first Key works
curator of experiments for the
Royal Society of London. In 1665 1665 Micrographia
he became Professor of Physics at 1674 An Attempt to Prove
Gresham College. the Motion of the Earth
Like many scientists of his 1676 A Description of
day, Hooke had a broad range of Helioscopes and Some
interests. His achievements Other Instruments
86
IF YOU DO NOT KNOW

THE NAMES OF THINGS,
THE KNOWLEDGE OF
THEM IS LOST
A SYSTEM FOR IDENTIFYING
ALL NATURE’S ORGANISMS
B
efore the 18th century, there characteristics, such as similarity
IN CONTEXT was no consistent naming of body parts, size, shape, and
system for animals and methods of getting food. Linnaeus
KEY FIGURE
plants. Botanists and zoologists also adopted a precise two-word
Carl Linnaeus (1707–78)
often did not know if they were (binomial) name for each species.
BEFORE discussing the same organism.
1682 John Ray, an English To overcome the problem, Swedish Early insights
botanist, proposes that the botanist Carl Linnaeus invented By 1730, while still a student,
plant kingdom be divided a revolutionary system, which is still Linnaeus began to have issues
into trees and two families in use today. He is known as the with the system for classifying
of herbaceous plants. “father of taxonomy”—the science of plants developed by Joseph Pitton
naming and classifying organisms. de Tournefort more than 30 years
1694 French botanist Joseph Linnaeus divided both the plant earlier. For Linnaeus, the
Pitton de Tournefort publishes and animal kingdoms into classes, characteristics of individual species
Eléments de Botanique. This orders, genera, and species. needed to be analyzed more closely
beautifully illustrated book Organisms were placed in these in order to produce a more thorough
becomes the botanical levels on the basis of shared taxonomic system.
classification benchmark
for half a century.
AFTER To work together over
Collaborative work is
1957 Sir Julian Huxley is the long distances, scientists
crucial for the advancement
first to use the term “clade” to need things to be
of scientific knowledge.
describe a common ancestor named with accuracy.
and all of its descendants.
1969 Robert Whittaker, an
American ecologist, argues for
a five-kingdom categorization
of life: Monera, Protista, Fungi, If you do not know
Misunderstandings
Plantae, and Animalia. the names of things, cause discrepancies in
the knowledge of scientific knowledge.
them is lost.

See also: Classification of living things 82–83 ■ Biological species concept 88–89
■ A modern view of diversity 90–91
been known by long impractical

names—for example, Plantago foliis
ovato-lanceolatis pubescentibus,
spica cylindrica, scapo tereti.
Linnaeus called this plant Plantago
In natural science, media, which was sufficient to
the principles of truth identify it. As well as being concise,
ought to be confirmed the Linnaean system describes
by observation. relationships between species.
Carl Linnaeus
Later developments Carl Linnaeus
Linnaeus constantly expanded
Systema Naturae; its 10th edition Born in rural southern Sweden,
(1758) became the starting point Linnaeus was educated at the
for modern animal classification. University of Uppsala, where
It was he who suggested that he began teaching botany in
In 1732, Linnaeus joined an humans were members of the 1730. He spent three years
expedition to Lapland, where he primate family. Much later, aided in the Netherlands, and, on
collected about 100 unidentified by Charles Darwin’s theory of returning to Sweden, he
species. These formed the basis of evolution by natural selection, divided his time between
his book Flora Lapponica, in which biologists accepted that a teaching, writing, and plant-
he aired his ideas about plant classification should reflect the collecting expeditions. At
Uppsala, 17 of his students
classifications for the first time. principle of common descent,
embarked on expeditions all
Three years later, Linnaeus which led to the methodology
over the world. Linnaeus was
wrote about his idea for a new known as cladistics. ■ a friend of Anders Celsius, the
hierarchical classification of plants inventor of the temperature
in a further book, Systema Naturae, scale. After his friend’s death,
Whales were once thought to be
and thereafter in arguably his fish, and were classified as such in Linnaeus reversed the scale
greatest work, Species Plantarum, an early edition of Linnaeus’s Systema so that freezing point was
published in 1753, which covered Naturae. Only later was it understood 32°F (0°C) and boiling point
7,300 species. Previously, plants had that they are actually mammals. 212°F (100°C). Linnaeus
has been described as the
“prince of botanists,” and the
philosopher Rousseau said of
him “I know no greater man
on Earth.” Linnaeus is buried
in Uppsala Cathedral; his
remains constitute the type
specimen—the specimen that
represents a species—used for
Homo sapiens.
Key works
1735 Systema Naturae

1737 Flora Lapponica
1751 Philosophia Botanica
1753 Species Plantarum
88
“REPRODUCTIVELY
ISOLATED” ARE THE
KEY WORDS
BIOLOGICAL SPECIES CONCEPT
IN CONTEXT
When two groups
KEY FIGURE “Species” can be of the same species
defined as population become reproductively
Ernst Mayr (1904–2005) groups that are able isolated they
BEFORE to reproduce. evolve separately.
1686 Naturalist John Ray
defines individual plant and
animal species as those that
derive from the same seed.
The capacity to Eventually, they
1859 Charles Darwin’s On the interbreed is key become separate
Origin of Species introduces to the definition species that cannot
the idea that species evolve of a species. mate with each other.
through natural selection.
AFTER
1976 The Selfish Gene by
B
Richard Dawkins popularizes y the early 20th century, it some point become separated by
gene-centered evolution: natural was accepted that multiple geography, mate choice, feeding
selection at a genetic level. species could evolve from a strategies, or other means, and then
1995 The Beak of the Finch common ancestor. However, it was begin to change through natural
not clear how this evolution process selection or genetic drift. Over time,
by Jonathan Weiner follows
actually occurred. In fact, there was as a result of this initial separation,
the work of biologists Peter
some debate about precisely what a two distinct species evolve, which
and Rosemary Grant on the
“species” was. In 1942, evolutionary cannot interbreed. This type
Galapagos Islands. biologist Ernst Mayr proposed a of speciation commonly occurs
2007 Massimo Pigliucci and new definition of species: groups in small populations of creatures
Gerd B. Müller use the term of interbreeding natural populations on remote islands.
“eco-evo-devo” to suggest how that are “reproductively isolated
ecology is among the factors from other such groups.” Key differences
affecting evolution. What this means is that two The biological species concept is
populations of the same species primarily focused on the breeding
living in the same area may at potential between organisms. Two

See also: Evolution by natural selection 24–31 ■ The role of DNA 34–37 ■ The selfish gene 38–39
■ Competitive exclusion principle 52–53
organisms may appear identical Sometimes, too, different animal

and live in the same place, but this species are able to mate and
does not mean that they are the produce offspring, as is the case
same species. For example, the of a female horse (Equus ferus
Western Meadowlark (Sturnella caballus) and a male donkey
neglecta) and Eastern Meadowlark Endless forms (Equus africanus asinus), which
(Sturnella magna) look similar and most beautiful and together can produce a hybrid—
have overlapping ranges, but they most wonderful have the mule. However, mules
have evolved to produce different been, and are themselves are generally incapable
songs. This prevents them from being, evolved. of reproduction, and therefore the
mating with each other, making Charles Darwin horse and donkey remain different
them two distinct species. species. Another example is the
Another scenario is when liger, a zoo-bred hybrid of a female
members of the same species look tiger and a male lion.
very different, but because they can Such anomalies highlight the
mate and reproduce they are still complexities of defining a species.
considered to be the same species. The biological species concept
The most obvious instance of this is of a species. Geographical remains the most popular, but
the domestic dog (Canis familiaris), separation alone does not prevent scientists are now looking at the
a species in which there are great species from reproducing if they idea of shared genes, and using
differences between individuals. are brought together. Evolutionary DNA sequence analysis. To date,
However, as is also evident, different divergences—such as the different no one has come up with a single
breeds are capable of reproduction mating songs of the Western definition that covers every known
with each other, and therefore and Eastern Meadowlarks—are species, and it seems unlikely
belong to the same species. what prevents interbreeding. that anyone ever will. In the
The biological species concept absence of better models, Ernst
Complex permutations is not applicable to asexual Mayr’s biological species concept
According to the biological species organisms, such as bacteria, provides an extremely useful
concept, the potential for inter- or asexual creatures—for example, way of thinking about species
breeding is key to the definition species of whiptail lizard. and evolution. ■
Alternative species concepts

Although Mayr’s idea about be based on genetics, such as
biological speciation is perhaps DNA or RNA base sequences, or
the most common way to define on phenotypes, such as the size
species and explain how they of certain body parts or
evolve, it is far from the only particular markings, such as the
one. In fact, there are more than arrangments of spots on insects’
20 recognized species concepts, wings. The evolutionary species
ranging across two broad groups: concept is based on species
typological and evolutionary lineages. A species is defined as
concepts. Typological species the organisms that share a
Male fireflies are an example of concepts are based on the idea lineage from the time when the
a typological species. They emit a that a population of individuals of species initially split off until
pattern of flashes to attract females, the same type—or sharing the extinction, or until an additional
who recognize their species’ code same set of traits—are what splitting off and creation of a
and flash back—if they wish to mate. makes up a species. The traits can new species.
90
ORGANISMS
CLEARLY CLUSTER
INTO SEVERAL
PRIMARY KINGDOMS
A MODERN VIEW OF DIVERSITY
B
efore biologists had the with simple nucleus-free cells),
IN CONTEXT equipment and techniques and eukaryotes (such as animals
needed to scrutinize the and plants with larger, more
KEY FIGURE
microscopic structure of living complex cells).
Carl Woese (1928–2012)
things, biological diversity was In the 1970s, the American
BEFORE split simply into animal-like and biologist Carl Woese claimed that
1758 Systema Naturae (10th plant-like organisms. Then, in the even this system failed to account
edition) by Carl Linnaeus 20th century, better microscopes for the diversity among microbes—
classifies known life into two began to reveal deeper differences the smallest living things. He
kingdoms: animals and plants. that could not be seen with the focused on ribosomes—minuscule
naked eye. By the 1960s, picking
1937 French biologist Edouard up on an idea first proposed by
Sulfur-dependent archaea
Chatton divides life into Edouard Chatton in the 1930s, the organisms thrive in the hot geothermal
prokaryotes (bacteria) and need for a new division of living pools of Yellowstone National Park,
eukaryotes (organisms with things emerged, placed between Wyoming, in conditions that would
complex cells). prokaryotes (such as bacteria, kill most other organisms.
1966 German biologist Willi

Hennig establishes a system
of classification based on
clades—groups of organisms
based on common ancestry.
1969 American ecologist
Robert Whittaker divides
life into five kingdoms:
bacteria, protists, fungi,
plants, and animals.
AFTER
2017 A consensus among
biologists accepts a seven-
kingdom classification of life.

See also: Early theories of evolution 20–21 ■ Evolution by natural selection 24–31
■ The role of DNA 34–35 ■ A system for identifying all nature’s organisms 86–87
Kingdom of their own
For most of the history of
biology, fungi were considered
Carl Woese’s three-domain tree to be plants. Even the great
classifier of organisms Carl
Linnaeus included them in his
Archaea kingdom Plantae. It was only
Thermoproteus with the invention of more
Methanococcales powerful microscopes that
Extreme Eukaryotes the differences in fungi began
Bacteria halophiles to be better understood. It is
Animals
Cyanobacteria now known that chitin, a
Plants
Bacteroides complex carbohydrate and
Fungi
Purple bacteria component of fungus cell
Protists
walls, is not found in plants.
Also, fungi make their food
by digesting rotted material,
According to Carl Woese, all organisms can be whereas plants make food by
separated into three main categories or “domains.” absorbing light energy in
These divisions are based on similarities in the photosynthesis.
ribosome structure found in the cells of the groups DNA analysis shows that
of organisms within each domain. fungi are far removed from
plants in the evolutionary
tree of life: they are, in fact,
grains that all cells need in order to A decade before Woese proposed genetically closer to the
make protein—and devised what he his theory, Robert H. Whittaker branch that gives rise to
called the “three-domain system.” had recognized animals, plants, animals. These same studies
This gave him a new perspective and fungi as separate eukaryotic show that certain aquatic
on the branches of Charles Darwin’s kingdoms, with all other eukaryotes molds—traditionally classified
evolutionary “tree of life.” Woese placed in the protist kingdom, as fungi—are not related to
found big differences in the and bacteria constituting a fungi, while some disease-
chemical makeup of ribosomes fifth kingdom. Whittaker’s protist causing microbes are fungi
among tiny microbes, with one kingdom covered eukaryotic that have evolved to become
microscopic parasites.
group as far from other prokaryotes organisms such as amoebas that
as bacteria are from humans. did not fit the other categories.
Some protists were closer to
Revising the tree of life animals, some closer to plants,
Woese’s third domain of organisms, and others not close to either.
known as archaea, is superficially They did not match the tree of life
similar to bacteria, but has some model, in which clades—groups
strange properties. Many thrive in of organisms with a common
extreme habitats. Some—uniquely ancestry—spring as branches
among living things—generate from the previous fork.
methane in oxygen-deprived places, Woese sought a classification
such as deep marine sediments, or system that reflected the intricacies
inside warm digestive cavities, of evolution—with main branches
such as those of belching, flatulent on the tree of life splitting into
plant-eating mammals. Other smaller ones, and even tinier twigs Fungi, such as this bright
archaea inhabit lakes that are ten that end in the leaves of individual yellow jelly fungus growing on a
fallen tree, are no longer classified
times saltier than seawater, or hot species. In the future, the complex as plants. Fungi are genetically
acidic pools fed by geothermal heat tree of life may reveal even more closer to animals.
that would kill anything else. evolutionary categories. ■
92
IN CONTEXT
SAVE THE
KEY FIGURE
Edward O. Wilson (1929–)
BIOSPHERE
BEFORE
1993 The UN proclaims
December 29 as the
International Day for
AND YOU MAY

Biological Diversity.
1996 The Song of the Dodo
by American science writer
SAVE THE WORLD

David Quammen explores
the nature of evolution and
extinction as habitats become
more and more fragmented.
HUMAN ACTIVITY AND BIODIVERSITY AFTER

2014 The Sixth Extinction
by environmental journalist
Elizabeth Kolbert shows how
humans are causing a sixth
mass extinction of species.
2016 In Half-Earth, Edward
Wilson proposes that Earth
can be saved by dedicating
half of it to nature.
B
iodiversity is the variety of
life on Earth—in all forms
and at every level, from
genes to microbes to humans and
all other species, including those
yet to be discovered. Humans rely
on biodiversity for food and fuel,
shelter, medicine, beauty, and
pleasure. For other species, it also
provides nutrients, seed dispersal,
pollination, and reproductive
success. No living thing could
survive without biodiversity.
Ecologists have identified
growing threats to biodiversity,
many of them driven by human
actions. The current rate of species
extinction is thought to be up to
1,000 times greater than it was

See also: Biodiversity hotspots 96–97 ■ Animal ecology 106–113 ■ Island biogeography 144–149 ■ Biodiversity and
ecosystem function 156–157 ■ Biomes 206–209 ■ Mass extinctions 218–223 ■ Deforestation 254–259 ■ Overfishing 266–269
The effects of human activity on biodiversity animals because they may no

longer be able to find places to feed
The five human activities or rest along their normal routes.
that most seriously affect Native species and ecosystems are
biodiversity on Earth can be
represented by HIPPO, the also disrupted by the introduction,
acronym devised by Edward accidentally or deliberately, of new
Wilson, with the relative species. These invasive species.
severity of each reflected can threaten the food supply or
in the order of the letters. other resources of native species,
1. Habitat destruction carry disease, and become a
predatory threat. The brown tree
snake, for example, was brought
accidentally to the island of Guam
on a cargo ship, and has caused
the extirpation (the extinction of a
species in a particular area) of 10 of
the island’s 11 native bird species.
5. Overharvesting 2. Invasive
by hunting species
or fishing Air and water poisoning
Any kind of pollution threatens
biodiversity, but air and water
pollution are particularly harmful.
Burning fossil fuels, for example,
releases the waste gases sulfur
dioxide and nitrogen oxide into
the air; these return as acid rain,
4. Human population 3. Pollution causing water and soil acidification
and affecting ecosystem health and
biodiversity. Ozone emissions at
before 1800, when humans began loss of habitats that once supported ground level can also damage cell
to dominate the planet. The first particular species. This destruction membranes on plants, curbing their
use of the term “biodiversity,” in can occur as a result of natural growth and development. ❯❯
1988, was by American biologist causes, such as fire or flood, or,
Edward O. Wilson, who became more commonly, through the
known as the “father of biodiversity.” expansion of agricultural land,
He later highlighted five key threats timber harvesting, and overgrazing
to biodiversity using the acronym by livestock. Deforestation, in
HIPPO: habitat destruction; particular, has contributed hugely It is that range of biodiversity
invasive species; pollution; human to habitat loss, with around half of that we must care for—the
population; and overharvesting by the world’s original forests now whole thing—rather than
hunting and fishing. cleared, mainly for agricultural use.
just one or two stars.
Some habitats are not destroyed
Habitat wreckers but rather broken up or divided into
David Attenborough
British broadcaster and naturalist
The Red List of the International more isolated units by human
Union for Conservation of Nature interventions, such as building
(IUCN) includes more than 25,000 dams or other water diversions.
threatened species. Of these, This habitat fragmentation is
85 percent are endangered by the particularly dangerous for migratory
94 HUMAN ACTIVITY AND BIODIVERSITY

Rapid population growth has
generated further damage to the
environment. The world’s human
population has risen from less
than 1 billion in 1800 to more than
We should preserve 7 billion, and is expected to reach
every scrap of biodiversity nearly 10 billion by 2050. As the
as priceless while we population grows, so do other
learn to use it and threats to biodiversity: increasing
come to understand numbers of invasive species are
what it means spread through trade and travel;
to humanity. urban development and resource
Edward O. Wilson extraction destroy habitats; more
pollution is created; and land is
overharvested. The impacts of
Edward O. Wilson
human population growth will be
Born in Alabama in 1929, difficult to limit, as ever more
Edward Osborne Wilson was people rely on food and shelter to
left blind in one eye after a survive, and demand ever more
fishing accident aged seven, Water pollution is caused mainly goods in an increasingly global
and switched interests from by sewage or by chemicals consumer society.
birdwatching to insects. He absorbed into water as it flows
discovered the first colony of off agricultural land. This pollution Upsetting the balance
fire ants in the US when he reduces oxygen levels in water, Population growth also drives
was only 13, and later making survival more difficult for overharvesting, the final human-
attended the University of some species, particularly when made threat to biodiversity in the
Alabama and Harvard. combined with water temperatures HIPPO acronym. Found in forestry,
Wilson’s work has focused
that have risen due to climate livestock grazing, and commercial
primarily on ants but also
extends to the study of change. Freshwater streams used agriculture, overharvesting can
isolated ecosystems, known by certain species of spawning also arise from targeted hunting,
as “island biogeography.” A fish, for example, can be made gathering, and fishing, as well as
leading environmentalist, uninhabitable by pollution. unintentional harvesting, such
he has spearheaded efforts Some organisms can absorb a as fish discarded from catches.
to preserve biodiversity and substance, such as an agricultural
educate people about it. He chemical, more quickly than they
has been awarded over 150 can excrete it, in a process known
prizes, including the National as bioaccumulation. Initial, low
Medal for Science, the Cosmos concentrations of chemicals may
Prize, and two Pulitzer Prizes not be a problem. However, as those
for nonfiction, and was named chemicals accumulate through the
one of the century’s leading food chain—from phytoplankton to
environmentalists by Time
fish to mammal, for example—they
and Audubon magazine.
can reach levels that cause birth
Key works defects and disrupt hormone levels
and immune systems.
1984 Biophilia
1998 Consilience: The Unity
of Knowledge Poaching, forest clearance, and
2014 The Meaning of Human other human activities have largely
Existence contributed to the status of the African
western lowland gorilla as a “critically
endangered” species.
THE ORDERING OF THE NATURAL WORLD 95

The building of railways across the
US was accompanied by hunters hired
to decimate the buffalo population that
had sustained Native American tribes.
By the end of the 19th century, only a
small number of wild buffalo survived.
against natural disasters and

human-made shocks, including
climate change, and provide
recreational, medicinal, and
biological resources.
Although the threats to
biodiversity from human activity
are serious, ways to protect it
are being developed. Foremost
is a “sustainable” approach to
harvesting and agriculture that
allows species—such as fish, trees,
When the rate of harvest exceeds pollution and climate change. More or crops—to be maintained at a
the rate of replenishment through than 80 per cent of the species on stable level and even increased
either reproduction or human the IUCN’s Red List are affected by over time. Official protected status
activities such as tree planting, more than one of the five major for areas of land, water, and ice
the harvest is not sustainable, biodiversity threats. can help sustain threatened
and without regulation could result Biodiversity maintains the species, while national and
in the extinction or extirpation health of the ecosystems of the international agreements and
of species. planet. Ecosystems are a delicate negotiations can mitigate the
A study of the IUCN’s Red List balance of living creatures, both impact of both legal and illegal
in 2016 showed that 72 per cent plant and animal, as well as the trade, such as poaching. Public
of species listed as threatened or soil, air, and water in which they education also helps people to
near-threatened are harvested at live. Healthy ecosystems provide better understand their potential
a rate that means their numbers resources that sustain human and impacts on biodiversity and how
cannot be balanced by natural all other life, improve resilience to protect it for future generations. ■
reproduction or regrowth. Some
62 per cent of species are at risk Anthropogenic biomes grouped into six main
from agricultural activity alone, categories: dense settlements;
such as livestock farming, tree The biosphere—all the areas of villages; croplands; rangeland;
felling, and the production of Earth and its atmosphere that forested; and wildlands.
crops for food, fuel, fibres, and contain living things—consists Unlike other biomes, which
animal fodder. of biomes, which are large can range across continents,
ecosystems based on a specific anthropogenic biomes are a
Protecting biodiversity environment, such as desert or mosaic of pockets over Earth’s
In reality, the five HIPPO threats tropical rainforest. The impact surface. According to ecologists,
identified by Wilson are interrelated, of human actions on biodiversity more than 75 per cent of Earth’s
and there is generally no single and the consequent reshaping ice-free land has been affected
of much of the planet have led by at least some form of human
reason why any particular species
ecologists to reassess biomes activity, particularly in dense
is endangered. Agricultural
and suggest that a designation settlements (urban areas), which
development, for example, can of anthropogenic (manmade) account for over half the world’s
not only destroy a habitat, but can biomes is now necessary. population, and villages (dense
also releases greenhouse gases into Anthropogenic biomes are agricultural settlements).
the atmosphere, contributing to air
96
WE ARE IN THE
OPENING PHASE OF
ABIODIVERSITY
MASSHOTSPOTS
EXTINCTION
A
biodiversity hotspot is an and increasing challenge of mass
IN CONTEXT area with an unusually extinctions of species caused by
high concentration of the destruction of premium habitats,
KEY FIGURE
animal and plant species. The term Myers argued that priorities had
Norman Myers (1934–)
was coined in 1988 by Norman to be set to establish where to
BEFORE Myers, a British conservationist, concentrate resources to conserve
1950 Theodosius Dobzhansky to describe areas that are both as many lifeforms as possible.
studies plant diversity in biologically rich and deeply
the tropics. threatened. Facing the huge Defining hotspots
Initially, Myers identified ten
AFTER hotspots crucial for conserving
2000 Myers and collaborators The lush hillsides and forests of
Arunachal Pradesh, India, are part plant species that were endemic
reevaluate the list of hotspots of the Indo-Burma biodiversity hotspot. (did not grow anywhere else on
and add several new ones, The area contains some 40 per cent Earth). By 2000, he had refined the
bringing the total to 25. of India’s animal and plant species. concept to focus attention on
2003 An article in American
Scientist criticizes the
concentration of conservation
effort on hotspots, saying that
this neglects less species-rich
but still important “coldspots”.
2011 A team of researchers
confirm the forests of east
Australia as the 35th hotspot.
2016 The North American
coastal plain is recognized
as meeting the criteria for a
global biodiversity hotspot—
and becomes the 36th.

See also: Human activity and biodiversity 92–95 ■ The ecosystem 134–137
■ Deforestation 254–259 ■ Sustainable Biosphere Initiative 322–323
looks like an antelope; it was seen

for the first time in 1992, in the
Annamite Mountains of Vietnam.
The endangered Irrawaddy dolphin
is found along the coastlines of
Our welfare is intimately Southeast Asia and the islands
tied up with the welfare of of Indonesia. Other rare animals
wildlife … by saving the lives include Eld’s deer, the fishing cat,
of wild species, we may be and the giant ibis.
saving our own.
Norman Myers Protective measures Norman Myers
Conservation agencies agree on
targets for every hotspot. They list Myers was born in 1934
species that are threatened and and grew up in the north of
make plans to conserve and manage England. He studied at the
those areas with suitable habitat University of Oxford before
and viable populations of target moving to Kenya, where he
regions that fulfilled two criteria: plants and animals. Sites are ranked worked as a government
the area must contain at least according to how vulnerable and administrator and teacher.
1,500 vascular plants (plants with irreplaceable they are. During the 1970s, Myers
roots, stems, and leaves) that were Myers’ two criteria have been studied at the University of
endemic, and it must have lost at criticized by those who say they do California, Berkeley, where his
interest in the environment
least 70 percent of its primary not take account of changing land
grew. He raised concerns
vegetation (the plants that originally use in regions where less than 70
about deforestation for cattle
grew in the area). Conservation percent of good habitat has been ranching, describing it as the
International, an environmental destroyed. The Amazon rain forest, “hamburger connection.”
agency that uses Myers’ concept to for example, is not within a hotspot Myers raised the concept
guide its efforts, now lists 36 such but the forest is being cleared faster of biodiversity hotspots in
regions. Although they represent than anywhere else on Earth. ■ the article “Threatened
only 2.3 percent of Earth’s land Biotas: ‘Hotspots’ in Tropical
surface, they are home to nearly Forests,” published in The
60 percent of the planet’s plant, Environmentalist in 1988.
amphibian, reptile, mammal, and In his first book, Ultimate
bird species—and a high Security: The Environmental
proportion of these species live only Basis of Political Stability, he
in their respective hotspot. We are into the opening argued that environmental
Most hotspots lie in the tropics stages of a human-caused problems lead to social and
biotic holocaust—a wholesale political crises. In 2007, Time
or subtropics. The one facing the
elimination of species—that magazine hailed Myers as a
highest threat level is the Indo- Hero of the Environment.
Burma area in Southeast Asia. could leave the planet
Only 5 percent of the original impoverished for at least
habitat remains, but its rivers, five million years. Key works
wetlands, and forests are vital for Norman Myers
1988 “Threatened Biotas:
the conservation of mammals,
Hotspots in Tropical Forests”
birds, freshwater turtles, and fish. 1993 Ultimate Security: The
Animals unique to this area Environmental Basis of
include the saola, a forest-dwelling Political Stability
mammal that is related to cattle but
THE VARI
OF LIFE
ETY
100 INTRODUCTION
Louis Pasteur reveals Charles Elton publishes

Dutch lens-makers Hans that wine’s fermentation Animal Ecology, which
and Zacharias Janssen process is caused by germs; sets out many of the
invent the compound his discovery sparks the fundamental principles
microscope. development of germ theory. of animal behavior.
1590 1866 1927
1676 1885
Antonie van Albert Frank coins the

Leeuwenhoek term “mycorrhizae,” in
identifies “animalcules,” reference to the symbiotic
opening up the field relationship between
of microbiology. fungi and tree roots.
O
ur understanding of the about them for many years. In trees with the fungi attached
variety, behavior, and the 1860s, French chemist Louis to their roots were healthier than
interraction of organisms Pasteur and German microbiologist those without. The fine filaments,
has advanced considerably since Robert Koch developed the germ or hyphae, of the fungi make the
Aristotle discovered that bee theory of disease, highlighting the roots more efficient at obtaining
colonies have a queen and workers. harmful role played by bacteria. nitrate and phosphate nutrients
Huge advances in technology, Subsequent research has also from the soil. In return, the fungi
field observations, and laboratory highlighted their positive roles: get sugar and carbon from the tree.
experiments have increased our facilitating digestion; inhibiting
knowledge, and the modern study the growth of other, pathogenetic Connected lives
of animal behavior—ethology— bacteria; “fixing” or converting No organism lives in isolation from
continues to throw up surprises. nitrogen into molecules that aid the rest of its ecosystem. The
plant growth; and breaking down behavioral interactions between
Life under the microscope dead organic material, which them are complex and much is still
Until the microscope was invented, releases nutrients for the food web. being discovered about them. One
no one knew that bacteria even Another discovery made of the greatest contributions in this
existed, let alone what they did. possible by microscopy was of the field was made by British zoologist
Bacteria were first observed by mutualistic relationship between Charles Elton, whose 1927 classic
Dutch microscopist Antonie van fungi and trees, published by Animal Ecology established many
Leeuwenhoek in 1676, using German plant pathologist Albert important principles of animal
an instrument he had built himself. Frank in 1885. Studying what he behavior, including food webs and
He called these tiny organisms first assumed was a pathological food chains, prey size, and the
“animalcules,” but little was known infection, Frank discovered that concept of ecological niches.
THE VARIETY OF LIFE 101
In Man Meets Insect Thermoregulation In the US, the Human

Dog, Konrad Lorenz by Bernd Heinrich Microbiome Project is
describes the instinctive explains how insects launched, to map all the
behavior of animals and are able to control their microbes associated with
its evolutionary origins. own temperature. a healthy human body.
1949 1981 2007
1947 1960 2005
David Lack publishes Jane Goodall sets up The first gene mapping
an article on variations a camp in Tanzania to confirms that humans
in bird clutch size as research chimpanzees in share 97–99 percent of
an evolutionary the wild, discovering many their DNA with the other
adaptation. traits shared by humans. great apes.
Ethology, which looks at animal Field observations are a key tool in a red spot on a parent’s beak when
behavior and its evolutionary basis ethological research. In the 1940s, they want food, will tap colored
and development, is a major British ornithologist David Lack marks painted on a model beak.
component in the modern study investigated the factors controlling
of organisms. Back in 1837, British the number of eggs birds laid Human traits
entomologist George Newport (clutch size). His food limitation As well as these short-term studies,
discovered that moths and bees hypothesis states that the number British primatologist and ethologist
could raise the temperature of their of eggs laid by a species has Jane Goodall conducted field
thorax by quivering their muscles. evolved to match the food available. observations over a longer period,
From the 1970s onward, German- Evolutionary pressure has created studying chimpanzees in Tanzania
American entomologist Bernd a correlation between clutch size from 1960 to 1975. Her findings
Heinrich and others uncovered and food availability. challenged the view that human
more thermoregulatory adaptations Austrian zoologist Konrad behavior is totally unique in the
that have helped insects thrive. Lorenz and Dutch biologist Nikolaas animal world, and indicated that
As heterotherms, they are able Tinbergen also studied animals in chimps are behaviorally closer
to maintain different temperatures the wild to help understand their to people than had generally been
in different parts of the body. behavior. Lorenz’s 1949 work Man assumed. She noted, for example,
Modern research now combines Meets Dog explains the loyalty that chimps display a whole range
laboratory experiments, field of a pet dog to its owner in terms of facial expressions and other body
observation, and new technology of canines’ instinctive loyalty language to indicate their mood, are
such as infrared thermography to their pack leader in the wild. toolmakers and users, often behave
to understand insect behavior Tinbergen’s field experiments cooperatively, and sometimes go
in ever more detail. showed how gull chicks, which tap into battle against rival groups. ■
102
IT IS THE MICROBES
THAT WILL HAVE THE
LAST WORD
MICROBIOLOGY
M
icrobes—bacteria, Scientists did not understand
IN CONTEXT molds, viruses, protozoa, microbes until they could see them.
and algae—are present The first observations began in the
KEY FIGURE
in every environment, living in soil, 17th century, using the recently
Louis Pasteur (1822–95)
water, and air. Some microbes invented microscope. These
BEFORE cause disease but most are vital for studies revealed a previously
1683 Dutch amateur scientist life on Earth. Among other things, unknown world teeming with
Antonie van Leeuwenhoek they break down organic matter so microbiotic life. Around the same
uses a microscope to observe that it can be recycled back into time, the word “germ,” originally
bacteria and protozoa. the ecosystem. meaning “seed,” was first used
Trillions of microbes also live on to describe these tiny organisms.
1796 Edward Jenner carries and in the human body. The most
out the first vaccination, using common of these microbes are Fighting disease
the cowpox virus to protect beneficial bacteria, which aid the Some 17th and 18th-century
against smallpox. digestion of food, produce vitamins, scientists believed that certain
and help the immune system find “germs” might cause diseases, but
AFTER and attack more harmful microbes. the prevailing view was that such
1926 American microbiologist maladies were the spontaneous
Thomas Rivers distinguishes result of inherent weakness in an
between viruses and bacteria. organism. It was not until the
1928 While studying painstaking laboratory work of the
influenza, Scottish 19th-century French chemist Louis
Pasteur that the “germ theory of
bacteriologist Alexander
Microbes are the worker bees disease” was proved.
Fleming discovers penicillin.
that perform most of the Pasteur began by looking at the
2007 An inventory of all important functions alcohol fermentation process. He
the microbes associated discovered that sourness in wine
in your body.
with a healthy human body was caused by external agents—
Dr Robynne Chutkan, microbes, or germs. A crisis in the
is completed. Microbiome expert and author
French silk industry, caused by an
epidemic among silkworms, then
allowed Pasteur to isolate and
identify the microorganisms that
caused the particular disease.

See also: Classification of living things 82–83 ■ The microbiological
environment 84–85 ■ The ecosystem 134–137
animal or human, was particularly

effective at enabling the body’s
immune system to fight off the
disease. At first, Pasteur faced
strong opposition and alarm at
Where observation the prospect, but he was able to
is concerned, develop vaccines for anthrax, fowl
chance favors only cholera, and rabies—the latter
the prepared mind. involving his first test on a human.
Louis Pasteur
Annihilating germs Louis Pasteur
The focus later shifted to finding
germ-killing agents, or antibiotics, Born in Dole, France, in 1822,
such as penicillin—discovered by Pasteur was the son of a poor
Alexander Fleming. A strategy of tanner. He was an average
annihilating microbes has been student, but he worked hard,
As he extended germ theory to followed ever since. Yet this obtaining his degree in 1842
human disease, Pasteur proposed “slash and burn” approach has and his doctorate in science
that germs invade the body and its drawbacks. It kills beneficial in 1847. After teaching in
cause specific disorders. Edward microbes as well as harmful ones, various universities, in 1867
Jenner, nearly 100 years before, and also promotes resistance in he became Professor of
had shown that a disease could be bacteria that can ultimately render Chemistry at the Sorbonne
in Paris. His major research
prevented with the application of antibiotics ineffective. ■
interest was the fermentation
a “vaccine”—a virus similar to that
process. Pasteur discovered
of the disease-causing microbe. that the fermentation of wine
The bacterium Enterococcus faecalis
Pasteur found that an attenuated, is a microbe found in the gut and bowel and beer was caused by
or weakened, form of a disease- of healthy humans. If it spreads to other germs—microbes. He also
causing germ, produced in a areas of the body, however, it can cause discovered that microbes
laboratory and injected into the host serious infections. could be killed by short, mild
heat treatment—a process
now named after him as
“pasteurization.” Pasteur’s
“germ theory” led to the wider
development of vaccines,
which remain a vital method
of disease control. In 1887,
he established the Pasteur
Institute, which opened in
1888 and continues to help
prevent and fight diseases.
Key works
1870 Studies on Silk Worm

Disease
1878 Microbes: Their Role in
Fermentation, Putrefaction,
and the Contagion
1886 Treatment of Rabies
104
CERTAIN TREE SPECIES

HAVE A SYMBIOSIS
WITH FUNGI
THE UBIQUITY OF MYCORRHIZAE
I
n 1885, a professor of plant
IN CONTEXT pathology at the Royal College
of Agriculture in Berlin named
KEY FIGURE
Albert Frank was the first to see a
Albert Frank (1839–1900)
connection between fungi growing
BEFORE on tree roots and the health of the
1840 German botanist trees. Frank realized that these were
Theodor Hartig discovers not pathological (disease-related)
a network of filaments on the infections but in fact underground
roots of pine trees. partnerships: far from suffering, the
trees seemed to benefit from better
1874 Hellmuth Bruchmann, nutrition. He invented a new term
a German biologist, notes for the partnership—“mycorrhiza,”
the “Hartig net” is made of from the Greek mykes, meaning
fungal filaments. fungus, and rhiza, meaning root.
AFTER Mycorrhizae in action Mycorrhizae on the root of a
1937 A.B. Hatch, an American False truffles are an example of soybean. In arbuscular mycorrhizae,
botanist, shows a beneficial the fungal side of this partnership. such as these, the tips of the hyphae
relationship between pine form clusters inside the plant’s root
Nineteenth-century Prussian cells, optimizing nutrient exchange.
trees and mycorrhizal fungus. botanists had found these fungi
1950 Swedish botanists Elias under spruce trees, and noticed
that each tree root was drawn absorb the soluble organic
Melin and Harald Nilsson
toward a truffle, and wrapped in compounds produced through a
show that plant roots can
a fungal husk. Although they did network of microscopic filaments—
extract more nutrients from the
not know it, the botanists were hyphae—called a mycelium.
soil with the aid of mycorrhizae. witnessing a phenomenon that Plants rely on root hairs to
1960 Another Swedish is vital to many ecosystems. absorb water and minerals, such as
botanist, Erik Björkman, shows Fungi are typically nourished nitrates and phosphates. But there is
that plants pass carbon into by a supply of organic matter, from a limit to how far plant roots can
mycorrhizal fungi in exchange which they extract food by external grow and therefore what quantity of
for phosphate and nitrate. digestion. A deep layer of forest nutrients the root hairs can absorb.
litter is perfect. They pour digestive The hyphae of mycorrhizae can
chemicals onto their meal and cover a much wider area, absorbing

See also: Evolution by natural selection 24–31 ■ Mutualisms 56–59
■ The ecosystem 134–137 ■ Energy flow through ecosystems 138–139
Mycorrhizae as
pollution indicators
Mycorrhizal fungi are not only
Beneficial exchange between good for the health of plants—
mycorrhizae and plant roots they can also act as indicators
of the health of the entire
Mycorrhiza Plant environment. Laboratory
Supplies sugar from
photosynthesis experiments with these fungi
have shown that some grow
badly in the presence of toxins,
Connects plants in an which means that they can be
extensive network used to detect pollutants in
the air or soil. For instance,
some fungi fail to grow when
Increases uptake of exposed to heavy metals such
The mutualistic as lead or cadmium, and
relationship between water and nutrients
because different kinds of
mycorrhizae and plants fungi react differently to
is highly evolved. As
Allows plants to share environmental change, certain
many as 90 percent of
all plant species rely on nutrients with others species can be used to identify
fungi for nutrients and specific kinds of pollution.
protection. In return, Mycorrhizae are also useful
plants supply the fungi Boosts protection indicators of the health of their
with a vital food source. against soil diseases native habitat. Many form
cauliflower-like growths
on tree roots, but these are
a much greater amount of minerals. fungi in this way. Trees supported smaller in polluted soil. The
When the fungal hyphae attach to by mycorrhizae are more resistant trees themselves may also
the plant roots, they extend the root to drought and disease, and can respond to pollution with
system, causing extra nutrients to even communicate alarm signals by weaker shoot growth, but
seep into the plant. releasing chemicals in response to the mycorrhizal response is
Albert Frank realized that this attack by herbivores. This fungal more acute and serves as a
partnership worked both ways. It network connecting trees has been valuable early-warning sign
was a winning combination for both dubbed “the wood-wide web.” ■ of a habitat in decline.
plant and fungus. In exchange for
passing on a share of its minerals,
the fungus receives sugar from the
plant—made by photosynthesis in
the leaves and transported to the
roots via the plant’s sap. This boosts
the nutrient supply that the fungus [the fungus] performs
derives from dead organic matter. a “wet nurse” function
and performs the
Ancient networks entire nourishment
Fossils of plants dating from 400 of the tree from
million years ago—when vegetation the soil.
was first spreading across dry Albert Frank
land—show traces of fungal threads. Weak growth in the russet
This suggests that the mycorrhizal brittlegill, a mycorrhizal fungus
of European and North American
partnership was key to the evolution spruce forests, can be an early
of terrestrial life. Today, the majority indicator of habitat air pollution.
of plant species continue to rely on
FOOD IS THE
BURNING
QUESTION
ANIMAL ECOLOGY
108 ANIMAL ECOLOGY
T
he concept of food chains— knowledge of individual plant and
IN CONTEXT the idea that all living animal species to the cells in a
things are linked through beehive—each “cell” of knowledge
KEY FIGURES
their dependence on other species is important in its own right, but
Charles Elton (1900–91),
for their food—dates back many by putting them all together
George Evelyn Hutchinson centuries, but it was not until the something much more than the
(1903–91) early 20th century that scientists sum of the parts is created—
BEFORE developed the concept of food the “beehive” of ecology.
Ninth century Arab writer chains forming a food web. Nowadays, the study of animal
Al-Jaziz introduces the The pioneer of this thinking was ecology focuses on how animals
concept of the food chain British zoologist Charles Elton, interact with their environment,
in Kitab al-Hayawan (Book whose book Animal Ecology (1927) the roles played by different
of Animals), concluding that describes what he called the “food species, why populations rise
cycle”. He later went on to develop and fall, why animal behaviour
“every weak animal devours
theories that encompassed more sometimes changes, and the
those weaker than itself”.
complex interactions between impact of environmental change on
1917 American biologist animals and the environment— animals. The principle underlying
Joseph Grinnell first describes insights that underpin modern the work of animal ecologists is
an ecological niche in his animal ecology. He likened our that there is generally a balance
paper, “The niche relationships
of the California Thrasher.” Food web
AFTER
1960 American ecologist and
philosopher Garrett Hardin
publishes an essay in the
magazine Science in which
he states that “every instance Whale
of apparent coexistence must Albatross
be accounted for.” Krill

Phytoplankton
1973 Australian ecologist
Robert May publishes Stability
and Complexity in Model Arrow worm Leopard
Ecosystems, in which he uses seal
mathematical modeling to Blue-green Radiolarians
demonstrate that complex bacteria
ecosystems do not necessarily Penguin
lead to stability. Schooling fish
Dolphin
Copepod
Squid
A food web is a graphic depiction

Killer whale
of the feeding connections between
different species within an ecological
community. This example illustrates Marine worm
Bottom-feeding
Seaweed fish
the relationships within a marine
ecosystem, in which killer whales are
the apex predators and phytoplankton
are the primary producers.

See also: Keystone species 60–65 ■ The food chain 132–133 ■ The ecosystem
134–137 ■ Energy flow through ecosystems 138–139 ■ Trophic cascades 140–143
Forecasting the effects
of climate change
in nature, so if the population Ecologists examine changes

of a given species grows too large to animal populations and
distribution and apply climate
it will be regulated, most often by a
change models to forecast how
lack of food. However, relationships these will change further in
between organisms and their the future, over the course of
environment change from place 5, 10, 50, or more years. In the
to place and through time. Arctic, for example, where
average temperatures are
Chain of dependence rising more rapidly than
In Animal Ecology, Elton outlined anywhere else, the sea ice is
the key principles of the study of contracting. As a result, polar
animal communities: food chains bears have to travel farther in
and webs, food size, and ecological search of ice where they can
niches. Each food chain and web, A spider traps a damselfly, catch seals, rest, and mate.
he asserted, is dependent on demonstrating that the principle The farther they swim, the
producers: plants and algae that of food size can be modified by the more energy they burn. As
comparative aggression and strength the sea ice declines, the polar
support plant-eating consumers of the predator and its prey. bears starve. Scientists
(herbivores). These herbivores in monitor their numbers and
turn support one or more levels of movements and compare this
meat-eating consumers (carnivores). against rivals of their own species data with changes in sea ice.
Large carnivores generally eat to ensure there is enough food for The polar bear plays a
smaller animals, but because small themselves and their offspring. vital part in the ecology of the
animals reproduce more quickly, Arctic. As an apex predator
their numbers are able to support Food size and keystone species, it must
the larger predators. One of Elton’s most important have access to seals, which
Competition for food is very tight points was the notion that food are its almost exclusive diet.
near the top of a food web. Although chains exist primarily because The number of seals regulates
apex (top) predators, such as big of the principle of food size. He the density of polar bears,
cats and large birds of prey, have no explained that every carnivorous while polar bear predation in
natural predators, this often means animal eats prey between upper turn regulates the density and
that they have to defend territories and lower limits. Predators are reproductive success of seals.
physically unable to catch and
consume other animals above a
certain size because they are not
large enough, strong enough, or
skillful enough. That is not to say
that predators cannot kill and eat
Every animal is closely larger animals than themselves; a
linked with a number of weasel can easily kill a larger rabbit
other animals living because it is more aggressive.
around it—and these are However, an adult lioness, one
largely food relations. of the world’s top predators, is not
Charles Elton capable of killing a healthy adult
African elephant. Likewise, A lone polar bear surveys the
a dragonfly larva on the bottom sea for prey from a piece of floating
ice in the Arctic. The shrinking
of a pond may be able to prey area of sea ice in the region
on a small tadpole, but it would threatens this species’ survival.
not be able to eat an adult frog. ❯❯
110 ANIMAL ECOLOGY

Snowshoe hare and The Sword-billed Hummingbird,
lynx population cycles a native of South America, has a long
bill, which enables it to suck up nectar
In Canada’s boreal forests, from the long flowers of Passiflora
the favored prey of lynx are mixta, a species of passionflower. As
snowshoe hares. Charles Elton it feeds, it spreads the plant’s pollen.
examined the relationship
between the populations
of these two species, using working in the early decades of the
data covering the period 20th century, an organism’s niche
1845–1925. When hares are was defined as its habitat. He
numerous, lynx hunt little studied birds called thrashers in
else. After their population California and observed how they
reaches its peak density, fed, nested, and hid from predators
the hares struggle to find in the dense undergrowth of the
enough plant food. Some chaparral shrubland. However, a
starve, while others are niche is more complex than simply
weakened and are more the place where an organism lives.
easily caught by predators, Oxpeckers and buffalo share exactly
including lynx, which feed the same habitat—open grassland –
very well for a time. When
but their requirements for survival
hare numbers continue to
are very different: the buffalo graze
fall, this affects the lynx.
They are forced to hunt on the grasses, while oxpeckers
less nutritious prey, such Animals may be capable of killing derive their food from the ticks they
as mice and grouse. much smaller prey, but it is simply peck from the buffalos’ hide.
As they struggle to find not worth the effort. Wolves hunt Charles Elton explored the
enough to eat, lynx produce medium-sized or large mammals concept of ecological niches in
smaller litters or even stop such as elk. If those mammals more depth. For him, food was
breeding altogether. Some disappear from their environment, the primary factor in defining an
starve to death. A decline in they find it hard to catch sufficient animal’s niche. What it ate and
the lynx population sets numbers of smaller animals such what it was eaten by were crucial.
in one or two years after the as mice to sustain them; the Depending on the habitat, a
hare population has bottomed energy they use finding small prey particular niche could be filled by
out, a cycle that repeats every is greater than the energy they a different animal. Elton cited the
eight to eleven years. gain by consuming them. example of a niche that was filled
Plants cannot run away or fight
back, so different considerations
apply to herbivores when it comes
to food size. There is a maximum
size of seed that a given finch,
for example, can fit in its bill, so
larger finches have an advantage Observation of species in the
over smaller species. Similarly, wild convinces me that the
individual species of hummingbirds existence and persistence of
can drink nectar only from flowers species is vitally bound up
up to a certain size, depending with environment.
on the length of their bill. Joseph Grinnell
A Canadian lynx captures a Ecological niches
snowshoe hare, its preferred prey.
When hares are plentiful, a lynx An animal or plant’s niche is its
will eat two every three days. ecological role or way of life. For
American zoologist Joseph Grinnell,

interacts with other organisms and example, koalas depend almost
with its environment. For example, entirely on eucalyptus leaves, and
each species of trout—and other Hyacinth Macaws in the Pantanal
fish—has its own range of water region of Brazil eat virtually nothing
salinity, acidity, and temperature but the hard fruits of two species of
Different species press that it can tolerate, as well as a palm trees—these are specialists.
against one another, like range of prey and river- or lake-bed Animals rarely occupy the whole
soap bubbles, crowding conditions. This makes some better of their niche width, owing to
and jostling, as one competitors than others, depending competition between species. Part
species acquires … some on the conditions of the habitat in of the habitat requirement of North
advantage over another. which they live. Seen as the father American bluebirds is dead trees
G. Evelyn Hutchinson of modern ecology, Hutchinson with old woodpecker holes in which
inspired other scientists to explore they lay their eggs and raise their
how competing animals use their young. Although suitable holes are
environment in different ways. common in many forests, bluebirds
An animal or plant’s niche cannot occupy all these holes
width comprises the whole range because they are often out-
of factors it requires to allow it to competed by more aggressive
by birds of prey that hunted small thrive. Brown rats, raccoons, and starlings. Therefore their realized
ground-dwelling animals such as starlings are examples of animals niche—the places they actually
mice and voles. In a European with a broad niche width in that occupy—is not as extensive as their
oakwood, that niche would be filled they are able to survive in a wide potential (or fundamental) niche.
by Tawny Owls, while on open variety of conditions. Such species Many animals share some
grassland Kestrels would fill the role. are called generalists. Other animals aspects of their niche, but not
Elton also argued that an animal have narrow requirements. For others. This is called niche ❯❯
could not only tolerate a certain set
of environmental conditions, but
could also change them. The tree-
felling and stream-damming
activity of beavers is one of the
most dramatic examples, creating
habitats for fish in dammed pools,
woodpeckers in dead trees, and
dragonflies around pool margins.
Niches and competition

British-born zoologist G. Evelyn
Hutchinson, working at Yale
University from the 1950s to the
1970s, examined all the physical,
chemical, and geological processes
at work in ecosystems and
proposed that any organism’s role
in its niche includes how it feeds,
reproduces, finds shelter, and
A true specialist, a koala bear

requires 2.5 lb (1kg) of eucalyptus
leaves a day. This species is found
in the wild only in Australia, where
eucalyptus is common.
112 ANIMAL ECOLOGY

Three major overlap. If different species live in
types of ecological the same habitat and have similar
pyramid lifestyles, they will be in
competition but they may be able
to live in close proximity if some
Pyramid
of numbers aspects of their behavior or diet The basic process in
1
Osprey
differ. This arrangement is known trophic dynamics is the
as niche partitioning. For example, transfer of energy from
10 various anole lizards on Puerto one part of the ecosystem
Northern pike Rico successfully occupy the same to another.
100 areas because they select perching Raymond Lindeman
Perch locations in different parts of trees.
1,000 There are limits to niche
Bleak overlap. When two animals with
identical niches live in the same
10,000 place, one will drive the other
Freshwater shrimp
to extinction. This concept—the
competitive exclusion principle— numbers to produce a pyramid
was outlined by Joseph Grinnell of biomass that represented the
Pyramid
of biomass in 1904 and developed in a paper amount of living matter in a given
Wolf published by Russian ecologist area at every level. This took
340 lb
per sq mile
Georgy Gause in 1934, becoming into account the fact that some
known as Gause’s law. organisms are much larger than
Red fox
1,700 lb others, but because it showed
per sq mile Pyramid of numbers comparative biomasses at a fixed
Snowshoe hare
Charles Elton used a pyramid as point in time, it produced anomalies.
17,000 lb per sq mile a way of graphically representing For example, in a pond, the mass
the different levels in a food chain, of the phytoplankton producer
Grass with the producers at the bottom, (microscopic organisms that are the
18 million lb per sq mile the primary consumers on the foundation of the aquatic food web)
level above, and so on. Often, the may not be as great as the mass of
primary consumers—insects, the fish consumers at a particular
in particular—will outnumber the point in time, so the pyramid
Pyramid
of energy Apex producers, but the higher levels will be inverted. However,
predators of consumers will become less phytoplankton reproduce quickly
0.01% numerous toward the top of the
Secondary pyramid. This system does not
carnivores
0.1% take account of parasites; fleas
Carnivores and ticks on mammals and birds
1% will far outnumber the total of all
the vertebrates in an ecosystem.
Herbivores
10% In 1938, German-born animal
ecologist Frederick Bodenheimer
Producers modified Elton’s pyramid of
100%
Ecological pyramids represent Microscopic organisms, including

quantifiable data in an ecosystem. these diatoms, form a significant part
Numbers show the population size of of all ecological pyramids. Their huge
individual species in a trophic level; numbers and rapid reproduction
biomass, their relative presence; and provide mass and energy for the
energy, who eats what and how much. species higher up the pyramid.
when conditions, such as sunlight herbivore, some of the energy Tench feed on snails, which graze
and nutrients, are right. Over time, transfers to the animal. When on periphyton—a mixture of microbial
the mass of the phytoplankton will a predator eats the herbivore, organisms that cling to plants. By
reducing the number of snails, tench
far outweigh that of the fish. it receives a smaller amount of that increase the periphyton biomass.
energy, and so on.
Trophic pyramids Published in 1942, Lindeman’s
American ecologist Raymond Ten Percent Law explains that above. Assessing energy transfer,
Lindeman proposed a pyramid of when organisms are consumed, however, requires a lot of information
energy, called the trophic pyramid, only about 10 percent of the energy about energy intake, as well as the
showing the rate at which energy transferred from them is stored as number and mass of organisms.
is transferred from one level to the flesh at the next trophic level. The
next as herbivores eat plants, and energy model creates a more Future thinking
predators eat herbivores. An realistic picture of the condition Relationships between organisms
organism’s trophic level is the of an ecosystem. For example, if and their environment change
position it occupies in a food chain. the biomass of weed and fish in from place to place and through
Plants and algae are at trophic level a pond is the same, but the weed time. Global climate change is one
1, herbivores at level 2, and the first reproduces twice as fast as the fish, example of environmental factors
level of predators is at 3. It is rare the energy of the weed would be that will increasingly affect animal
for there to be more than five levels. shown to be twice as large. Also, communities. Some changes have
Plants convert the sun’s energy there are no inverted pyramids— already taken place, but one of the
into stored carbon compounds, there is always more energy in the challenges of ecological thinking
and when a plant is eaten by a lowest trophic level than the one in the future is to forecast others. ■
114
BIRDS LAY THE NUMBER

OF EGGS THAT PRODUCE
THE OPTIMUM NUMBER
OF OFFSPRING
CLUTCH CONTROL
W
hy do some birds lay eggs are removed from a nest; the
IN CONTEXT more eggs than others? bird will re-lay repeatedly to
For example, Blue Tits compensate for the loss.
KEY FIGURE
lay nine eggs, Northern Flickers six, Instead, Lack said, the number
David Lack (1910–73)
and Robins four. In the 1940s, of eggs laid by any species has
BEFORE British ornithologist and evolutionary evolved to fit with the food supply
1930 British geneticist ecologist David Lack proposed an available. In other words, nature
Ronald Fisher combines explanation that rapidly gained favors clutch sizes that correspond
Gregor Mendel’s work on support. He argued that the clutch to the maximum number of young
genetics with Charles size (number of eggs laid) was not
Darwin’s theory of natural controlled by the female’s ability to
Blue Tit nests contain an average of
selection, and argues that the lay eggs, since birds can lay many nine eggs, although the females can lay
effort spent on reproduction more eggs than they typically do. many more. David Lack proposed that
This fact can be demonstrated by the clutch size is determined by the
must be worth the cost.
replacement experiments, in which likely amount of available food.
AFTER
1948 David Lack extends his
theory of optimal clutch size
in birds to include litter size
in mammals.
1954 Lack develops his food
limitation hypothesis further
in The Natural Regulation
of Animal Numbers, to
encompass birds, mammals,
and some insect species.
1982 Tore Slagsvold proposes
the nest predation hypothesis,
which states that clutch size
is related to the likelihood of
the nest being attacked.

See also: Animal ecology 106–113 Animal behavior 116–117 ■ The food chain
132–133 ■ The ecosystem 134–137
■
■ Ecological resilience 150–151

Siblicide and the
Blue-footed Booby
that the parents are likely to be able Blue-footed Boobies are

to sustain. So, if a pair of birds can seabirds native to the Pacific
Ocean. They get their food
only find enough food to feed six
from the ocean, but come
chicks, but the female has laid 12 to rocky shores and cliffs to
eggs, those young will be hungry breed. The female lays two
and may starve. If she has laid just Laying a clutch which eggs, roughly five days
one egg, although the chick will will result in a smaller apart, so that by the time the
be raised successfully, most of brood than … could be fed second chick hatches, the first
the available food will have been and reared successfully … one has already grown
unused. So neither the 12-egg nor confers advantages. considerably. When food is
the one-egg scenarios are good Tore Slagsvold plentiful, the parents can find
reproductive strategies; instead, enough to feed both offspring
laying six eggs offers the best until they fly the nest (fledge).
chance of raising the most offspring. However, when food is scarce,
This theory became known as the larger chick will peck its
the food limitation hypothesis, or junior sibling to death. The
older chick can then get more
Lack’s principle, and it was later
food, and is more likely to
generalized by him and others to of food during the long day-length
fledge. If it does not murder
cover litter size in mammals and of summer compared with the its sibling when food is scarce,
clutch size in fish and invertebrates. shorter day-length in the tropics. both chicks may starve.
However, other factors may This behavior, based
The “latitude trend” also apply. Higher mortality rates exclusively on the availability
Lack’s hypothesis also suggested in high latitudes—where winters of food, is called “facultative
an answer to another puzzle: why are harsh—may have led to the siblicide.” In contrast, masked
most bird species have bigger evolution of large clutch sizes. This boobies practice “obligate
clutches at higher latitudes. On is because the chances of survival siblicide”—the first-hatched
average, birds near the equator lay until the next breeding season chick nearly always kills its
about half the number of eggs laid are low, and the reduced population brother or sister, regardless
by the same species in the far results in more food being available of how much food is available.
north. This “latitude trend” could be for the survivors next season.
explained by a greater availability In 1982, Tore Slagsvold, a
Norwegian evolutionary ecologist,
advanced the nest predation
hypothesis, which proposes that
high rates of nest predation result
in smaller clutches. If a nest with
many chicks is found by a predator,
Clutch size increases more work by the parent birds will
with increasing latitude have been wasted than if the nest
and day length because … contained fewer chicks. Also,
a longer day enables the parents raising a large clutch are
parents to find more food. more likely to be seen by predators,
David Lack because of the extra activity. Some
ecologists have argued that the Blue-footed Boobies are driven
relative abundance of predators in to siblicide by genetic factors.
The murder of a sibling can benefit
the tropics has been more important the perpetrator while also ensuring
than food supply in the evolution of the survival of the entire species.
small clutch sizes at low latitudes. ■
116
THE BOND WITH A TRUE

DOG IS AS LASTING AS
THE TIES OF THIS EARTH
CAN EVER BE
ANIMAL BEHAVIOR
A
ny dog owner will describe
IN CONTEXT the companionable and
loyal relationship they
KEY FIGURES
enjoy with their pet. The Austrian
Konrad Lorenz (1903–89),
zoologist Konrad Lorenz set out
Nikolaas Tinbergen
to explain this behavior in Man
(1907–88) Meets Dog (1949). He described the
BEFORE behavior of dogs and other pets as
1872 Charles Darwin’s The substantially innate, “instinctive
Expression of the Emotions in activity,” as opposed to behavior
Man and Animals posits that learned through conditioning.
behavior is instinctive and has Lorenz proposed that such hard- Ducklings imprinting is an example
wired behavior helped the animal of instinctive behavior that can be
a genetic basis. manipulated—to make them imprint
survive as a species. For example, a
on humans or even inanimate objects.
1951 Nikolaas Tinbergen’s domestic dog’s loyalty to its human
The Study of Instinct lays master originates in the natural
down the foundations and behavior of its wild ancestors, when devising rigorous field
theory behind ethology, the which were loyal to the pack leader experiments that could be repeated,
study of animal behavior. because this had benefits in terms so that the findings could be
of hunting success and safety. recognized as facts, not anecdotes.
AFTER The term “ethology” was coined
1967 Desmond Morris, Field experiments by American entomologist William
a British zoologist, brings Lorenz was not alone in his theories. Morton Wheeler in 1902 to describe
ethology to bear on human Other biologists working in the field the scientific study of animal
behavior in his popular book included fellow Austrian Karl von behavior. Ethologists study animals
The Naked Ape. Frisch and Dutch biologist Nikolaas in their natural habitats, combining
Tinbergen, who studied animals laboratory studies and fieldwork
1976 British evolutionary in their natural environments. Until in order to describe an animal’s
biologist Richard Dawkins then, most animal behavior studies behavior in relation to its ecology,
publishes The Selfish Gene, had taken place in laboratories or evolution, and genetics.
describing how most of an artificial settings, so the behavior Ethologists found that in certain
animal’s behavior is designed witnessed was not entirely natural. situations, an animal will have a
to pass on its genes. Studying animals in the wild had predictable behavioral response.
its own challenges, particularly They called this a “fixed action

See also: The selfish gene 38–39 ■ Field experiments 54–55 ■ Keystone species 60–65 ■ Animal ecology 106–113 ■ Clutch
control 114–115 ■ Using animal models to understand human behavior 118–125 ■ Thermoregulation in insects 126–127
pattern” (FAP). A FAP has set Four elements of ethological experimentation

characteristics. It is species-
specific; it is repeated in the same
way every time and is not affected
by experience. The triggers for the Development
Causation
behavior (“sign stimuli”) are highly What stage is the animal
What triggered
specific and may involve a color, in its life cycle, and does the
the behavior in
pattern, or sound. For example, male behavior change as
the first place?
sticklebacks respond aggressively the animal develops?
when another male enters their
streambed patch. Ethologists
suggest this is triggered by seeing
the male’s red underbelly.
Nikolaas Tinbergen found When studying
that some an artificial sign stimuli animal behavior,
work better than the real thing. He scientists consider
investigated the begging behavior
these four elements
of herring gull chicks, which peck
at a red spot on the parent gull’s
beak to make it regurgitate food.
He found that chicks will also peck
at a model of the gull’s beak, yet
when they were offered a narrow Function
Evolution How does the behavior
red pencil with three white lines How is the behavior
at the end, the chicks pecked at increase the animal’s
related to the animal’s chance of survival or
this even more enthusiastically. evolution or ancestry?
Tinbergen called this a reproductive success?
“supernormal stimulus,” showing
that instinctive animal behavior
can be manipulated artificially. ■
Konrad Lorenz Born in Vienna, Austria, Lorenz and other birds, as well as
was enthralled by animals from mammals, is instinctive and
an early age and kept fish, birds, occurs shortly after birth. Lorenz
cats, and dogs. The son of an demonstrated the theory by
orthopedic surgeon, he studied quacking like a duck at newly
medicine at Vienna University, hatched ducklings. He soon
graduating in 1928, and gained had a flock of ducklings that
his Ph.D. in zoology in 1933. His followed him everywhere.
numerous pets became the first
subjects of his studies. Lorenz is Key works
perhaps best known for describing
the phenomenon known as 1952 King Solomon’s Ring:
“imprinting.” This is when a New Light on Animal Ways
newly hatched chick bonds with 1949 Man Meets Dog
the first thing it sees (usually its 1963 On Aggression
parent) and will follow it around. 1981 The Foundations for
The behavior, seen in ducks Ethology
REDEFINE “TOOL,”
REDEFINE “MAN,” OR ACCEPT
CHIMPANZEES
AS HUMANS
USING ANIMAL MODELS TO UNDERSTAND
HUMAN BEHAVIOR
120 USING ANIMAL MODELS TO UNDERSTAND HUMAN BEHAVIOR

The Primates Tree
IN CONTEXT
Humans and their
KEY FIGURE closest relatives,
Jane Goodall (1934–) chimpanzees, are both
primates. This shows
BEFORE 750,000–550,000 ya how primates have
1758 Carl Linnaeus, the father evolved over the last
66 million years.
of taxonomy, dares to classify 6 mya
humans within the rest of
nature, calling us Homo
sapiens (“wise man”).
Great
1859 Charles Darwin’s apes
theory of evolution further
challenges the established Lesser
apes
view that man is different
from the animal kingdom. Old world
monkeys
AFTER
1963 Konrad Lorenz publishes New world
monkeys
On Aggression, proposing 35 mya
that warlike behavior in
humans is innate. Primates
40 mya
1967 Desmond Morris,
a British zoologist and
ethologist, publishes The 65 mya
Naked Ape: A Zoologist’s
Study of the Human Animal,
a major study that describes Modern Chimpanzee/
human Neanderthal Gorilla Bonobo Orangutan
human behavior in the
context of the animal kingdom.
Gibbon Monkey Baboon Tarsier Lemur Loris
M
odern molecular studies scientific community remained
mapping the genomes convinced that humankind was
of humans and other different from the rest of nature.
animals have confirmed a theory It was largely the work of British
that was first suggested by Charles primatologist Jane Goodall that
Darwin in the mid-19th century— opened our eyes to the similarities
that we share a common ancestor between chimps and man. In 1961,
with the great apes. Today, few in an excited communication
In reality, we are Pan narrans, scientists would dispute that the to her mentor, Louis Leakey,
the storytelling chimpanzee. common chimpanzee (Pan Goodall announced an observation
Terry Pratchett troglodytes) and the bonobo or that would shake the scientific
British fantasy author pygmy chimpanzee (Pan paniscus) establishment: she had seen
are our closest living relatives. The a chimp using a tool. It was the
study of these animals therefore first time this behavior had
offers us a unique chance to learn been documented and it would
about ourselves and the origins of challenge perceived ideas of what
our behavior. Yet for many years the it means to be human.

See also: Evolution by natural selection 24–31 ■ A system for identifying all nature’s organisms 86–87 ■ Animal ecology
106–113 ■ Animal behavior 116–117
Goodall’s knowledge of natural

history had impressed Leakey
on their first meeting in 1957 and
he offered her a job studying the
behavior of chimpanzees. As an
anthropologist and paleontologist,
Leakey believed in evolutionary
theory, which proposed that humans
and the great apes—chimpanzees,
bonobos, gorillas, and orangutans—
in the family Hominidae (Great
Apes), share a common ancestor.
Making connections
Leakey’s fieldwork focused on
looking for the “missing link”—
fossils of transitional forms
between that common ancestor Until this point, the scientific and A chimp uses a twig stripped of its
and humans. Chimpanzees had not popular consensus was that the leaves—a modified “tool”—to catch
been studied seriously in the wild ability to devise and make tools termites for consumption. Goodall first
recorded the ability of chimpanzees to
and such a study, he reasoned, marked humans out as superior
invent simple technologies in Gombe.
could throw light on the evolution to the rest of the animal kingdom.
of early humans. Goodall, a keen Goodall’s findings forced scientists
observer and free of academic ties, to think again. true unfettered behavior, Goodall
was the ideal choice for the work. Goodall’s camp was in Gombe was one of the first people to work
As Leakey had hoped, she provided Stream National Park, Tanzania, in the field of ethology, whereby
a fresh perspective on the theory where she studied a chimp biologists monitor animals in their
and was brave enough to say that community on the eastern shore natural environments and try to
chimps and humans were more of Lake Tanganyika. In choosing to understand their natural behaviors.
alike than had been imagined. live among chimps to witness their In her first few months at the ❯❯
Jane Goodall Born in London in 1934, Jane transformed our understanding

Goodall’s first meeting with a of chimpanzees and challenged
chimp was a stuffed animal that perceived ideas of our own place
her father named Jubilee. She was in the natural world. In 1965, she
interested in animal behavior from earned a Ph.D. in ethology from
an early age—once, she hid in a Cambridge University. Her many
henhouse for hours so that awards include France’s Legion
she could watch a chicken lay of Honor, given to her in 2006.
an egg. She left school at 18 and
worked in various jobs, before Key works
going to Kenya in 1957 and
meeting paleoanthropologist 1969 My Friends the Wild
Louis Leakey. With his support, Chimpanzees
in 1960 Goodall set up a research 1986 The Chimpanzees of
base in Gombe, Tanzania, where Gombe: Patterns of Behavior
she was to study chimpanzees 2009 Hope for Animals and
until 1975. Her work radically Their World
camp, the chimps fled from her, and then using them in termite Jane Goodall working, notebook in
but they then began to forget she mounds; the chimps were not only hand, at Gombe National Park in 2006.
was there. using tools but making them. The pioneering primatologist continues
her lifelong commitment to protect
Goodall sat for many hours endangered chimpanzees.
observing the chimps, keeping her Chimp technology
distance and quietly making field Goodall went on to witness nine
notes. One morning in November different tools being used by instead of numbers, suggesting
1961, she noticed a chimpanzee she chimps in the Gombe community. that her fieldwork was less than
called David Greybeard sitting over At the time, scientists questioned rigorous. Since then, however, many
a termite mound. He was poking Goodall’s methods and ridiculed other studies around the world have
blades of grass into the mound, her for giving the chimps names corroborated her findings: chimps in
pulling them out, and then putting the Congo have been observed
them into his mouth. She watched stripping twigs to use in termite
for some time before the chimp mounds; chimps in Gabon have
moved off. On reaching the spot been seen heading into the forest
where the chimp had been sitting, with a five-piece “toolkit” that
Goodall saw discarded grass stems included a heavy stick for opening
lying on the ground. Picking one up I viewed my bee hives and pieces of bark for
and poking it into the mound, she fellow man not scooping up the honey. In Senegal,
found that the agitated termites bit as a fallen angel, hunting parties of chimps have been
onto the stem. She realized the but as a risen ape. observed traveling with sticks that
chimp had been “fishing” for Desmond Morris they chew to a sharp point and use
termites with the grass stems, and British Zoologist like spears to kill bush babies.
transferring them into his mouth.
From talks with Leakey, Goodall More alike than different
knew this was a major discovery. Ethologists take behaviors studied
She also saw chimps modifying thin across several species to formulate
twigs by stripping them of leaves generalizations that apply to many

it is worth noting that these Chromosomal
percentages are based on genes evidence
that instruct the body how to make
proteins, which make up a very A strong piece of evidence
small part of the human genome in favor of a shared common
ancestor is seen by comparing
We admit that we (about 2 percent). It is likely that the
chromosomes. Chimpanzees
are like apes, but things that make humans different (and gorillas) have 24 pairs of
we seldom realize from chimpanzees can be found in chromosomes. Humans have
that we are apes. the regions of DNA called “junk only 23. Evolutionary scientists
Richard Dawkins DNA” because they were previously believe that when we diverged
British evolutionary biologist thought to be redundant. It is now from a common ancestor, two
understood that this junk DNA chromosomes in humans
holds vital information about how fused and this is why we have
and when genes are expressed. one less pair than other apes.
Still, the similarities between the On the ends of every
DNA of humans and the great apes chromosome, there are genetic
are striking. markers—or sequences of
species. The idea that animal DNA—called telomeres. In the
behavior could be a model for Meat-eating hunters middle of each chromosome
there is a different sequence,
human behavior took root in the During her studies, Goodall also
known as a centromere. If
work of ethologists in the 1950s and witnessed chimps eating meat two chromosomes have fused,
’60s, such as Konrad Lorenz, and hunting. As with tool-making, it should be possible to see
Nikolaas Tinbergen, and Karl von the idea that chimpanzees were telomere-like regions in the
Frisch. Studying animals in their carnivorous predators went against middle of the chromosome as
natural habitats, they saw how all received knowledge. At first, well as at each end. Also, the
complex the lives of animals were. scientists claimed it was aberrant fused chromosome would have
They began to understand social behavior, but as the research two centromeres. Scientists
interactions arising from instinct continued and more sightings were looked and found just this.
as well as learned behaviors. The made, it became established fact. Human chromosome 2 appears
animal studies held a mirror up Meat-eating has been reported in to be the fusion of chimp
to human behaviors. just about every area where chimps chromosomes 2a and 2b. It is
The persistent belief that have been studied, from Gombe almost beyond doubt that we
humans are totally different from and Mahale Mountains National share a common ancestor with
other species was firmly rebutted Park, Tanzania, to Tai National chimps, bonobos, and gorillas.
with the advent of gene mapping. Park, in Côte d’Ivoire. ❯❯
When the chimpanzee genome was
mapped in 2005—followed by the
other great apes—and compared
with the human genome, the
results were clear. Humans share
98.8 percent of their DNA with
chimps, 98.4 percent with gorillas,
and 97 percent with orangutans.
Humans and great apes are more
alike than they are different. Yet
An alpha male’s body language

says “keep away” to begging chimps
wanting a share of his prey in Gombe
National Park. The main source of prey
for chimps is the colobus monkey.
Conservation of chimpanzees
According to the Jane Goodall chimp communities when roads
Institute in Tanzania, the number are built through their territories.
of chimpanzees living in the wild Roads also encourage another
has plummeted over the last damaging activity—hunting
century. In 1900, there were an for bushmeat, a highly valued
estimated 1 million chimpanzees meat in Africa that includes
in Africa; today, there are fewer great apes. Roads enable
than 300,000. Habitat loss due to hunters from towns to travel
a rising human population in need directly into the bush. The
Orphaned chimps—their mothers of more space has had a huge protection of chimps focuses
killed for bushmeat—walk along a impact, as have industries such on land conservation and on
mud track with their keeper at a as logging and mining, which raising awareness both locally
conservation center in West Africa. destroy habitat and fragment and across the globe.
Such behavior has implications Although chimps sometimes hunt for the war was unclear; some
for human evolution. Science has alone, hunting tends to be a group researchers blamed the feeding
long questioned why and when activity. Chimps rampage through stations Goodall had set up in the
humans first began eating meat. the forest, coordinating their area, which may have encouraged
From prehistoric stone tools and positions and surrounding their unnatural congregations of chimps.
marks on bones, paleontologists prey. After the hunt, the food is The answer to the mystery came
know that the early hominids shared. This shows how early in March 2018, when a research
were using stone tools to cut meat ancestors of humans may have team at Duke and Arizona State
from animals bones 2.5 million developed cooperative behavior, Universities, US, digitized Goodall’s
years ago, but it is not known a factor that may have contributed meticulous check sheets and field
what they were eating between to their evolutionary success. notes from 1967 to 1972 and fed
then and 7 million years ago, them into a computer in order to
when the common ancestor Chimp warfare analyze the social networks and
of chimpanzees and humans A shocking revelation that came alliances of all the male chimps.
is thought to have lived. out of the Gombe camp was that Their findings revealed that the
It is likely that these early chimps are capable of violence, fracture in the community occurred
hominids hunted prey. Although murder, and in particular warfare— two years before the war broke
they did not have large canine once believed to be the preserve out, when an alpha male Goodall
teeth like chimpanzees, these are of humans. Between 1974 and 1978, called Humphrey took over the
not necessary for hunting and Jane Goodall watched as her troupe, alienating two other high-
killing small prey. Biologists have peaceful community of chimps
observed that chimps hunting fractured into two rival groups that
colobus monkeys grab them from then waged savage war on each
the trees and then kill them by other. Goodall was deeply upset
repeatedly thumping the bodies about the chimps’ activity, which
on the ground; early hominids included ambushes, kidnappings,
could have hunted and killed and bloody murder. The trigger
in a similar fashion long before
the earliest known tools.
Chimps may fight over territory
in order to acquire more resources
Cooperative behavior or mates, but some primatologists
Another aspect of chimps’ hunting maintains that such aggression is
behavior that is similar to that unnatural and provoked by human
of humans is the social element. impact on their habitat.

associated with food resources and piece of fruit hanging over one room
meat eating. When food is limited, (B). The bonobo in room A could
the chimps become more violent release the fruit but not get to it
in order to obtain the resources himself. The researchers found that
they need. Chimps are known to this bonobo would consistently
I’m determined my eat more meat when fruit is scarce. release the fruit, so that the other
great grandchildren one could reach it, helping a
will be able to Kissing cousins stranger, with no reward for himself.
go to Africa and The link between food scarcity Researchers also observed how
find wild great apes. and aggression in the common the sight of an unknown bonobo
Jane Goodall chimpanzee may explain why our yawning in a film would trigger
other evolutionary cousin in the a yawning response in bonobos
primate world, the bonobo (pygmy watching the film, suggesting a
chimp), is so peace-loving. These capacity for empathy. Other studies
small, placid chimps are omnivores have shown how bonobos comfort
but live in an environment where each other when in distress. Unlike
fruit is plentiful most of the time. the “negative” behavior that
ranking males called Charlie and They forage in groups, and tend humans share with chimps,
Hugh and causing them to split to use sex to relieve tensions in these traits mirror more laudable
off with some other chimps to the social situations. Conflict is rare human characteristics, such as
south. The two groups became in bonobo societies, which are compassion. Understanding such
more and more separate, feeding also matriarchal, unlike the male- behavior in bonobos could shed
in different parts of the forest. At dominated chimp communities. light on how our human social
first there was the odd aggressive An experiment carried out by behavior developed. ■
skirmish and then war broke out. researchers at Duke University,
Over four years, Humphrey and his North Carolina, in 2017 showed
Bonobos are very social primates.
cohorts killed every male in the that bonobos are also altruistic. Two Their capacity for empathy makes
southern group and took over their bonobos (unknown to each other) them less aggressive and may align
territory, as well as three surviving were put in adjacent rooms (A and B) them more closely with their human
females. It is thought that the full- with a fence between them and a cousins than the common chimpanzee.
blown war may have been due
to a lack of mature females in the
northern group. Power struggles
and fighting over a female all sound
very human.
Fights over resources

The long-running war witnessed
by Goodall is the only sustained
conflict among chimpanzees to
have been fully documented,
but violence within chimp
communities has been recorded
many times. Chimps have been
observed stealing and killing baby
chimps and rounding on a disliked
alpha male. In communities studied
in Uganda, males routinely beat
the females they mate with. It is
thought that this violent streak
running through chimps may be
126
ALL BODILY ACTIVITY

DEPENDS ON
TEMPERATURE
THERMOREGULATION IN INSECTS
I
nsects are usually described as known that many insects are
IN CONTEXT “cold-blooded,” or ectotherms. heterotherms, maintaining different
Unlike mammals and other temperatures in different parts of the
KEY FIGURE
“warm-blooded” endotherms, body, and are sometimes far warmer
Bernd Heinrich (1940–)
animals that maintain their body than the ambient temperature.
BEFORE temperature at a more or less
1837 In the UK, George constant level, insects have a The right temperature
Newport observes that flying variable body temperature that The main challenge facing insects
insects are capable of raising changes with their environment. is how to get warm enough to fly
their body temperature above In the early 19th century, but cool enough not to overheat.
the ambient temperature. however, British entomologist German–American entomologist
George Newport discovered that Bernd Heinrich explained in 1974
1941 Danish researchers some moths and bees raise the how moths, bees, and beetles could
August Krogh and Eric temperature of their thorax (the continue to function by controlling
Zeuthen conclude that the central part of the body, to which their own temperature. He realized
temperature of an insect’s wings and limbs attach) above that that insects’ thermal adaptations
flight muscles just before of the surrounding air by rapidly do not differ as much from those of
takeoff determine the muscles’ flexing their muscles. It is now vertebrates as had been thought.
rate of work during flight. Most flying insects have higher
metabolic rates than other animals
AFTER but their small body size means they
1991 German biologist Harald lose heat rapidly, so they cannot keep
Esch describes how muscle their temperature constant at all
“warm-up” plays a role in brood times. The minimum temperature
incubation and colony defense In insects… the active that allows an insect to fly varies
as well as flight preparation. flight muscles… are, from species to species, but the
metabolically, the most maximum temperature falls within
2012 Using infrared active tissues known. 104–113°F (40–45°C). To prevent
thermography, Spanish Bernd Heinrich overheating, insects can transfer
zoologist Jose R. Verdu shows heat from the thorax to the abdomen.
how some dung beetle species Many larger flying insects would
heat or cool their thorax to remain grounded if they were not
improve flight performance. able to increase the temperature of
their flight muscles. These insects

See also: Evolution by natural selection 24–31 ■ Ecophysiology 72–73
■ Animal ecology 106–113 ■ Organisms and their environment 166
Heat regulation
Honeybees are renowned for
controlling the temperature
of their hive. When it gets too
hot, they ventilate it by using
their wings to fan the hot air
out of the nest. When it gets
too cold, the bees generate
metabolic heat by rapidly
contracting and relaxing their
flight muscles. They also use
heat as a defense mechanism.
Japanese giant hornets are
fierce predators of honeybees.
Capable of killing large
numbers quickly, they pose a
serious threat to bees’ nests.
Hornets begin their attacks by
picking off single honeybees
at the entrance to the hive.
However, Japanese honeybees
defend themselves with
self-generated heat. If a hornet
attacks, they swarm around it,
“quiver” the muscles that control A tortoiseshell butterfly feeds on vibrating their wings to raise
the upbeat and downbeat of the a dandelion. Most butterflies can angle their collective temperature.
wings to generate heat before their wings upward in an attempt Since the hornet cannot
to cool down, in a process called
taking off. Once flying, the muscles tolerate a temperature above
behavioral thermoregulation.
use large amounts of chemical 114.8°F (46°C) whereas the
energy but only some of it is used bees can survive at almost
to beat the wings; the rest becomes Other insects use even more 118.4°F (48°C), the attacker
more heat. This, combined with the remarkable methods to regulate their eventually dies.
warmth of direct sunlight, means body temperature. When a mosquito
a flying insect risks overheating. drinks the warm blood of a mammal,
To solve this problem, many this raises its body temperature. To
species have a heat-exchange compensate, it produces droplets of
mechanism that shifts excess heat fluid that are kept at the end of the
from the thorax to the abdomen, abdomen; evaporative cooling of
allowing the insect to maintain these droplets lowers the insect’s
a steady temperature in its thorax. temperature. Dung beetles construct
balls of dung in which females lay
Range of techniques their eggs. Some dung beetles are
By changing the angle of their able to raise the temperature of their
wings, butterflies control their body thorax so they can roll heavier balls.
temperature. When they are trying The range of thermoregulation
to warm up, holding their wings techniques shows how life forms
wide open maximizes the amount evolve to better fit their environment.
of sunshine falling on them. When They can also inspire technology: This Japanese giant hornet is
they are trying to cool down, they arrays of solar panels angled to raiding nursery cells in a bees’
nest in Hase Valley, Japan. Hornets
move into shade or angle their track the Sun capture maximum seek to devour the bee larvae
wings upward so that less direct amounts of solar radiation—just like inside the cells.
sunlight shines on their surface. butterfly wings. ■
ECOSYST
EMS
130 INTRODUCTION
Nelson Hairston,
Richard Bradley Charles Elton Frederick Smith, and
describes how plants, develops the idea of The journal Ecology Lawrence Slobodkin’s
pollinating insects, the food web in Animal posthumously publishes “green world hypothesis”
and insectivores rely Ecology and introduces Raymond Lindeman’s article argues that the predator–
on one another in a the concept of the “The trophic-dynamic prey balance is key to
food chain. ecological niche. aspect of ecology.” flourishing ecosystems.
1718 1927 1942 1960
1859 1935 1957
Charles Darwin Arthur G. Tansley coins the G. Evelyn Hutchinson

describes food term ecosystem, arguing establishes the concept
webs in his On the that an environment and all of niche breadth at
Origin of Species. its living organisms have the Cold Spring
to be seen as a single, Harbor Symposia on
interactive whole. Quantitative Biology.
W
hen Aristotle wrote The concept of the ecosystem (“a They identified both the top-down
about plant and animal recognizable self-contained entity”) pressures exerted by predators and
species existing for the followed soon after, when in 1935, the bottom-up pressures exerted by
sake of others, he showed a basic British botanist Arthur Tansley limitations on food supply. Twenty
understanding of food chains— wrote that organisms and their years later, American ecologist
as have countless observers of the environment should be considered Robert Paine wrote of the trophic
natural world since ancient Greek one physical system. In his Ph.D. cascade effect—the way a system
times. Arab scholar Al-Jahiz thesis, American ecologist is changed by the removal of a key
described a three-level food chain Raymond Lindeman expanded species. He described changes to
in the 9th century, as did the on Tansley’s work, positing that the food web after the experimental
Dutch microscopist Antonie van ecosystems are composed of removal of the ocher starfish from
Leeuwenhoek in 1717. British physical, chemical, and biological an intertidal zone. This predatory
naturalist Richard Bradley processes “active within a space– echinoderm was shown to be a
published more detailed findings time unit of any magnitude.” keystone species, playing a crucial
on food chains in 1718, and in 1859, Lindeman also conceived the role within its ecosystem.
Charles Darwin described a “web idea of feeding levels, or trophic
of complex relations” in the natural levels, each of which is dependent Island isolation
environment in his book On the on the preceding one for its survival. Habitat fragmentation is now a
Origin of Species. The concept of a In 1960, the American team of major problem in most terrestrial
food web, with many predator-prey Nelson Hairston, Frederick Smith, environments because it leaves
interactions, was then further and Lawrence Slobodkin published specialist organisms isolated.
developed by Charles Elton in his findings on the factors controlling For that reason, research into the
classic Animal Ecology (1927). animals on different trophic levels. biogeography of islands—those
ECOSYSTEMS 131
Hal Caswell proposes Scientists at the

a “neutral” theory Biodiversity and
John Maynard Smith of biodiversity, suggesting Ecosystem Function
defines his that competitors are often conference in Paris
Evolutionarily Stable equal, and chance plays examine how the
Strategy (ESS) theory the deciding role in loss of species
in On Evolution. what does or doesn’t thrive. impacts ecosystems.
1972 1976 2000
1967 1973 1980 2015
Robert MacArthur Crawford Stanley Robert Paine coins the term A study of grassland plants
examines the Holling uses the “trophic cascade” after his suggests that biodiversity
biodiversity of term ecological field experiments show the increases an ecosystem’s
isolated communities resilience to show how effect on an ecosystem resistance during
in The Theory of ecological systems when a keystone species and resilience after
Island Biogeography. survive change. is removed. climate events.
surrounded by ocean but also evolutionarily stable strategy (ESS) after a trauma. Ecologists now
“islands” of distinct habitat to describe the best behavioral understand that ecosystems can
surrounded by a very different strategy for an animal competing have more than one stable state,
environment—is so important with others living in its vicinity. and that resilient systems are not
in ecology. In the US in the 1960s, This strategy depends on how the always good for biodiversity.
Edward O. Wilson and Robert other animals behave and is When the populations of many
MacArthur discovered key factors determined by the animal’s genetic species are declining or becoming
determining species diversity, success—if it makes the wrong locally extinct, ecologists are once
immigration, and extinction on decisions, it will not live long and more focusing their attention on
islands. James Brown later did cannot pass on its genes. The ecosystem resilience. Many,
similar work on animal populations overall balance between the including French ecologist Michel
in isolated patches of forest ridge in evolutionarily stable strategies of Loreau, believe that if diversity in
California. Such work has showed all the animals in an ecosystem is an ecosystem is reduced, the whole
how to identify species most at risk called the evolutionarily stable state. system will be less likely to resist
of extinction due to isolation. Canadian ecologist Crawford major impacts such as the effects of
Stanley Holling introduced the idea climate change. Today, Loreau and
Stability and resilience of resilience—how ecosystems others are working toward finding a
One major contribution to the persist in the wake of disruptive new general theory that can
understanding of ecosystem changes such as fire, flood, or explain the relationship between
dynamics was the concept of the deforestation. A system’s resilience ecosystem biodiversity and
evolutionarily stable state. In the can be seen in its capacity to absorb resilience in order to understand
1970s, British biologist John disturbance, or the time it takes to and combat the effects of today’s
Maynard Smith used the term return to a state of equilibrium environmental challenges. ■
132
EVERY DISTINCT PART

OF NATURE’S WORKS
IS NECESSARY FOR THE
SUPPORT OF THE REST
THE FOOD CHAIN
The food chain

A
ll animals must eat other
IN CONTEXT living things in order to
receive the nutrients they
KEY FIGURE
Apex predator need to grow and function. A food
Richard Bradley (1688–1732)
chain shows the feeding hierarchy
BEFORE of different animals in a habitat.
9th century Arab scholar For example, the chain would show
Al-Jahiz describes a three- that foxes eat rabbits but rabbits
level food chain of plant matter, never eat foxes. Although there
rats, snakes, and birds. were earlier notions of a hierarchy
Larger predator of animals linked to each other in
1717 Dutch scientist Antonie (tertiary consumer) a food chain, British naturalist
van Leeuwenhoek observes Richard Bradley brought more
how haddock eat shrimp and detail to this idea in his book New
cod eat haddock. Improvements in Planting and
Gardening (1718). He noted that
AFTER each plant had its own particular
1749 Swedish taxonomist Carl Carnivore set of insects that lived off it and
Linnaeus introduces the idea (secondary consumer) proposed that the insects in turn
of competition. received the attentions of other
1768 John Bruckner, a Dutch organisms of “lesser rank” that fed
naturalist, introduces the idea on them. In this way, he believed
that all animals relied upon each
of food webs.
other in a self-perpetuating chain.
1859 Charles Darwin writes Herbivore
(primary consumer) Producers and consumers
about food webs in On the
Origin of Species. The modern concept of a food chain
explains that some organisms
1927 British zoologist Charles produce their own food. These are
Elton’s Animal Ecology outlines known as producers, or autotrophs.
principles of animal behavior, Plants and most algae fall into this
including food chains. Producer (autotroph) category, normally using the energy
of sunlight to convert water and
carbon dioxide into glucose, at the
ECOSYSTEMS 133
See also: Predator–prey equations 44–49 ■ Mutualisms 56–59 ■ Keystone species 60–65 ■ Optimal foraging theory 66–67
■ Animal ecology 106–113 ■ The ecosystem 134–137 ■ Trophic cascades 140–143 ■ Ecological resilience 150–151
Each species has a

specific place in nature,
in geographic location
and in the food chain.
Carl Linnaeus
same time as releasing oxygen. of the food chain, but there is always An apex predator, such as the bronze
This process, photosynthesis, is the a producer at the bottom, and all whaler shark, has no natural predators.
first step towards creating food. In levels above it are consumers. In the temperate waters of the ocean
off South Africa the shark can find vast
places where there is no sunlight, Animals that only eat plants are quantities of sardines to eat.
organisms producing their own herbivores, or primary consumers,
food are called chemotrophs. Those and they include cattle, rabbits,
in the deep ocean, for example, get butterflies, and elephants. Those of prey. The animals at the top of
the energy they need from chemicals that eat only other animals are their food chain are apex predators.
released by hydrothermal vents. carnivores, or secondary consumers; They include consumers such as
Animals that eat producers and these include thrushes, dragonflies, tigers, killer whales, and golden
creatures that eat other animals are and hedgehogs. In turn, secondary eagles that are not preyed upon
called consumers, or heterotrophs. consumers may be eaten by larger by other animals.
There may be two, three, or more predators, or tertiary consumers, The food chain does not break
levels of these in any particular part such as foxes, small cats, and birds when plants and animals die.
Detritus feeders (detritivores) prey
Richard Bradley infections were caused by tiny on the remains, recycling nutrients
organisms, visible only with a and energy for the next generation
Born around 1688, noted British microscope. His investigations of producers to use.
botanist Richard Bradley gained into the productivity of rabbit
patrons after writing a Treatise warrens and fish lakes led to Food webs
of Succulent Plants at the age of his theories about predator–prey Observers after Bradley suggested
22. With no university education, relations. Bradley died in 1732. that animals were not simply part
he was nonetheless elected a of a food chain, but a larger and
Fellow of the Royal Society and more complex “food web” that
later became the first professor Key works comprises all the food chains in a
of botany at Cambridge. location. This idea was put forward
Bradley’s research interests 1716–27 The History of by Dutch naturalist John Bruckner
were wide-ranging, including Succulent Plants
in 1768, and later taken up by
fungal spore germination and 1718 New Improvements in
plant pollination. In some cases, Planting and Gardening Charles Darwin, who called the
Bradley was ahead of his time; 1721 A Philosophical Account of variety of connected feeding
he argued, for example, that the Works of Nature relationships between species
a “web of complex relations”. ■
134
IN CONTEXT
ALL ORGANISMS KEY FIGURE

Arthur Tansley
ARE POTENTIAL
(1871–1955)
BEFORE
1864 George Perkins Marsh,
SOURCES OF an American conservationist,

publishes Man and Nature,
FOOD FOR OTHER

which hints at the concept
of ecosystems.
1875 Austrian geologist
ORGANISMS
Eduard Suess proposes
the term “biosphere.”
THE ECOSYSTEM AFTER

1953 American ecologists
Howard and Eugene Odum
develop a “systems approach”
to studying the flow of energy
through ecosystems.
Paul Sears highlights
the role of ecosystems
in recycling nutrients.
1970 Paul Ehrlich and Rosa
Weigert warn of potentially
destructive human
interference in ecosystems.
B
ritish biologist Arthur
Tansley was the first to
insist that communities
of organisms in a particular area
had to be seen in a wider context,
including the nonliving elements
of that area. Tansley argued that
in a given region, all the living
organisms and their geophysical
environment together form a single,
interactive entity. Borrowing a
concept from engineering, he saw
the network of interactions as a
dynamic, physical system. On the
suggestion of his colleague Arthur
Clapham, he coined the word
“ecosystem” to describe it.
ECOSYSTEMS 135
See also: Animal ecology 106–113 ■ The food chain 132–133 ■ Energy flow through ecosystems 138–139 ■ The biosphere
204–205 ■ The Gaia hypothesis 214–217 ■ Environmental feedback loops 224–225 ■ Ecosystem services 328–329
Tropical coral reefs are some of the

most diverse ecosystems of all, full of
fish, sea turtles, crustaceans, mollusks,
and sponges, as well as corals.
This idea had been developing

long before Arthur Tansley
published his influential paper
on the subject in 1935. As early
as 1864, conservationist George
Perkins Marsh, in his book Man
and Nature, had identified “the
woods,” “the waters,” and “the
sands” as different types of habitat.
He examined how the relationship
between them and the animals and
plants that lived in them could be
upset by human activity.
Interconnected systems
By the 20th century, the idea had
taken hold that these and other
environments could be understood a single unit, and using the term nitrogen, and soil minerals, which
as discrete entities, with distinctive “biome” to describe the whole are essential to the functioning of
interactions between the living complex of organisms inhabiting the systems as a whole. The biotic
and inert elements within them. In a given region. components within an ecosystem
1916, American ecologist Frederic Tansley envisaged ecosystems not only interact with one another,
Clements built on this idea in his as being made up of biotic (living) but also with the abiotic parts.
work on plant succession, referring elements and abiotic (nonliving) Thus, within any given ecosystem,
to a “community” of vegetation as elements such as energy, water, the organisms adapt to both the ❯❯
Arthur G. Tansley A free-thinking Fabian socialist He retired in 1937, but

and atheist, Arthur Tansley was maintained a special interest
one of the most influential in conservation. Tansley was
ecologists of the 20th century. appointed the first Chairman of
Born in London in 1871, he studied the UK’s Nature Conservancy in
biology at University College 1950, five years before his death.
London, where he later taught.
In 1902, he founded the journal Key works
New Phytologist and he later
established the British Ecological 1922 Types of British Vegetation
Society, becoming founding editor 1922 Elements of Plant Ecology
of its Journal of Ecology. In 1923, 1923 Practical Plant Ecology
he took a break from teaching to 1935 “The use and abuse of
study psychology with Sigmund vegetational terms and
Freud in Vienna. He was later concepts,” Ecology
Sherardian Professor of Botany 1939 The British Islands and
at the University of Oxford. Their Vegetation
136 THE ECOSYSTEM

biological and physical elements matter: plants absorb carbon
of the environment. The different dioxide (CO2) from the atmosphere
types of ecosystem can be defined and nutrients from the soil to grow.
by their physical environments. These plants release life-sustaining
There are four categories of oxygen into the atmosphere by
ecosystem: terrestrial, freshwater, respiration, and provide food for
marine, and atmospheric. These animals. Animal excreta and dead
can be further subdivided into matter also release carbon, and
various types according to different provide material to be decomposed
physical environments and the by bacteria and fungi, in turn
biodiversity within them. Terrestrial providing soil nutrients for plants.
ecosystems, for example, can be Arthur Tansley also argued
subdivided into deserts, forests, that these internal processes
A small glacial lake, or tarn, in the
grasslands, taigas, and tundras. within an ecosystem conform to English Lake District. Each tarn has
what he described as “the great an ecosystem that varies according
Dynamic feedback universal law of equilibrium.” to many factors, including the degree
Tansley’s most important insight Self-regulating, these processes of nutrient enrichment in the water.
was that these discrete communities have a natural tendency toward
of living and nonliving components stability. The cycles within an Each ecosystem is located in a
form dynamic systems. In a ecosystem contain feedback loops particular area, with characteristics
terrestrial ecosystem, for example, that correct any fluctuations from unique to its environment, and
the organisms interact to recycle a state of equilibrium. behaves as a self-contained and
self-regulating system. Together,
the patchwork of ecosystems
The dynamic transfer of energy across the globe form what
Austrian scientist Eduard Suess
In this ecosystem, plants use the Sun’s energy
for photosynthesis. As shown by the pale arrows,
called the biosphere—the sum
this energy is passed on—to herbivores, who eat total of all ecosystems.
plants; to predators, who eat herbivores; and to
saphrophytes, who take energy from decomposing External factors
remains and transfer nutrients to the soil. Various external factors, such as
At each stage, some energy is lost as heat. climate and the geological makeup
Heat of the surrounding environment,
can affect an ecosystem. One
Heat constant external force that affects
Plants all ecosystems is the Sun. The
(producers) Heat supply of energy that it provides
enables photosynthesis and the
capture of CO2 from the atmosphere;
Frog some of this energy is distributed
(predator) through the ecosystem and through
the food chain. In the process, some
of this energy dissipates as heat.
Other external factors, however,
Fish
(herbivores) can arise unexpectedly to create
Nutrients
Saphrophytes pressures on ecosystems. All
and parasites ecosystems are subjected to
(decomposers)
external disturbances from time
to time, and must then go through
a process of recovery. These
disturbances include storms,
ECOSYSTEMS 137
more fragile, and if they are wholesale destruction of habitat
disturbed may never be able and the consequent depletion of its
to regain their equilibrium. biodiversity. In addition, some have
The resistance and resilience suggested that human influence
of an ecosystem is generally has created a new category
There is no waste in thought to be related to its of ecological systems, dubbed
functioning … ecosystems. biodiversity. If, for example, “techno-ecosystems.” For example,
All organisms, dead or alive, there is only one species of plant “cooling ponds” are manmade
are potential sources of food performing a particular function ponds, built to cool down nuclear
for other organisms. in the system, and that species is power plants, but they have become
G. Tyler Miller not frost-resistant, an abnormally ecosystems for aquatic organisms.
Science writer severe winter could deplete the The relationship between
species enough to have a major humans and natural ecosystems
impact on the system as a whole. is not all negative. In recent years,
In contrast, if there are several scientific data has fueled public
species with that role in the awareness of the benefits that
system, it is more likely that one ecosystems afford humankind,
will be resistant to the disturbance. including the provision of food,
earthquakes, floods, droughts, and water, nutrients, and clean air,
other natural phenomena, but are The human factor as well as the management of
increasingly the result of human Some disturbances can be severe disease and even climate. There
activity—through the destruction enough to be catastrophic for an is now a growing commitment from
of natural habitats by deforestation, ecosystem, damaging it beyond many governments across the
urbanization, pollution, and the the point of recovery and so world to use these benefits both
cumulative effects of anthropogenic causing a permanent change responsibly and sustainably. ■
(human-induced) climate change. in its identity, or even its demise.
Humans can also be responsible The fear is that much of the
The Eden Project, in Cornwall, UK,
for the introduction of invasive disturbance caused by human simulates a rain forest ecosystem in
species. Without these external activity has the potential to one of its giant dome greenhouses. The
factors, an ecosystem would cause such permanent damage, domes’ panels are slanted to absorb
maintain its state of equilibrium, particularly when it involves the plenty of light and thermal energy.
and retain a stable identity.
Resistance and resilience

Ecosystems are often strong
enough to withstand some natural
external disturbances and retain
their equilibrium. Some are more
resistant to disturbance than
others, and have adapted to the
particular disturbances normally
associated with their environment.
In some forest ecosystems, for
example, the periodic fires caused
by electrical storms cause only a
minor imbalance in the ecosystem.
Even when severely disrupted
by external disturbances, some
ecosystems have a resilience
that enables them to recover.
However, other ecosystems are
138
LIFE IS SUPPORTED
BY A VAST NETWORK
OF PROCESSES
ENERGY FLOW THROUGH ECOSYSTEMS
I
n 1941, Raymond Lindeman his Ph.D., but his paper was initially
IN CONTEXT submitted the final chapter of rejected by Ecology, for being
his Ph.D. thesis for publication too theoretical.
KEY FIGURE
in the prestigious journal Ecology. Lindeman had painstakingly
Raymond Lindeman
Titled “The Trophic-dynamic sampled everything in the lake,
(1915–42)
Aspect of Ecology,” it was about from the aquatic plants and
BEFORE the relationship between food microscopic algal plankton to the
1913 American zoologist chains and the changes over time worms, insects, crustaceans, and
Victor Shelford produces the in a community of species. fish that fed upon one another and
first illustrated food webs. Lindeman had spent five years depended on each other for their
studying the life forms in an aging existence. He stressed that the
1920 Frederic Clements lake at Cedar Creek Bog, Minnesota, community of organisms could not
describes how groups of plant and was especially interested in be properly understood on its own;
species are associated in the changes in the lake as, year instead, it must be examined in the
communities. by year, aquatic habitat gradually context of its wider surroundings.
gave way to land. He received The living (biotic) organisms and
1926 Russian geochemist
Vladimir Vernadsky sees that
chemicals are recycled between
living and nonliving things. Producers (plants and Primary consumers
1935 Arthur Tansley develops algae) depend on energy are dependent on an
gathered from the Sun and abundance of plants
the concept of the ecosystem. nutrients from decomposed and algae to eat.
AFTER organic matter.
Eugene Odum uses radioactive
elements to map food chains.
1962 Rachel Carson draws
Secondary consumers
attention to the accumulation Life is supported rely on an abundance of
of pesticides in food chains, by a vast network primary consumers as
in her book Silent Spring. of processes. their food source.
ECOSYSTEMS 139
See also: Ecological niches 50–51 ■ Nonconsumptive effects of predators
on their prey 76–77 ■ The food chain 132–133 ■ The ecosystem 134–137
Measuring
productivity
Boneworms are deep-sea creatures Lindeman’s trophic-dynamic
that feed on the remains of animals theory helped to clarify the
such as whales. They grow “roots” to idea of ecosystem productivity,
break down the bones, thereby recycling which ecologists had previously
nutrients from the dead material. defined in rather vague terms.
The productivity of a plant
or animal is measured by its
Lindeman also demonstrated how growth in organic material,
some of the energy at each trophic or biomass. This is never equal
level is lost as waste, or converted to the organism’s energy
into heat when organisms respire. input, because the conversion
By combining the results of his own of solar energy into leaf in the
study with data from a wide range case of plants, or the conversion
of other sources, he was able to of food into flesh in the case of
build a picture of this system as it an animal, is never 100 percent
worked in Cedar Creek Bog. efficient. Some energy is
British ecologist G. Evelyn released as heat, most of which
is lost via respiration—an
Hutchinson, considered to be one
essential aspect of metabolism
of the founding fathers of modern
in all living things.
ecology, was Lindeman’s mentor at Warm-blooded animals lose
the nonliving (abiotic) components Yale University. He recognized the a lot of heat when their body
(air, water, soil minerals) were importance of his student’s work to temperature is much higher
linked together by nutrient cycles the future development of ecology, than that of their surroundings.
and energy flows. This entire and he lobbied for Lindeman’s All animals also lose energy
system—the ecosystem—was paper to be accepted. Lindeman, when they excrete urine. In
the central ecological unit. who had always suffered from ill addition, not all the material
health, died in 1942 from cirrhosis in an animal’s food can be
Producers and consumers of the liver at the tragically young digested in its gut, and the
Lindeman’s research showed how age of 27, just four months before material that is expelled as
an ecosystem is powered by a his trophic-dynamic paper—now feces represents unused
stream of energy from one organism seen as a classic in its field—was chemical energy.
to another. The organisms can be finally published. ■
grouped into discrete “trophic levels”
(feeding levels)—from producers
(plants and algae), which absorb
energy in the form of sunlight to
make food, to consumers (animals).
“Primary consumers” are the
herbivores that eat the plants; … biological communities
“secondary consumers” are animals could be expressed as
that eat the herbivores. Each trophic networks or channels through
level depends on the preceding one which energy is flowing and
for its survival. At the same time, being dissipated…
dead material accumulating from G. Evelyn Hutchinson
each stage is broken down by This thermal image of an
decomposers, such as bacteria and elephant shows how some of the
animal’s heat is lost. Both its body
fungi, and materials in the form of temperature and its manure are
nutrients are recycled back to feed warmer than the surroundings.
plants and algae.
140
IN CONTEXT
THE WORLD
KEY FIGURE
Nelson Hairston (1917–2008)
BEFORE
IS GREEN
1949 Aldo Leopold publishes
A Sand County Almanac,
drawing attention to the
ecological impact of hunting
wolves on mountain plant life.
TROPHIC CASCADES AFTER

1961 Lawrence Slobodkin,
an American marine
ecologist, publishes The
Growth and Regulation of
Animal Populations, a key
ecology textbook.
1980 Robert Paine describes
the “trophic cascade effect,”
when predators are removed
from an intertidal ecosystem.
1995 The reintroduction of
gray wolves to Yellowstone
National Park sets in motion
a series of ecosystem changes.
S
oon after the end of World
War II, Aldo Leopold, an
ecologist and one of the top
wildlife management experts in the
United States, challenged the view
that wolves should be eradicated
because they threatened livestock.
In A Sand County Almanac, he
wrote of the destructive effect that
removing this top predator would
have on the rest of the ecosystem.
In particular, he said, it would lead
to overgrazing of mountainsides by
deer. Leopold’s view was an early
expression of the idea of trophic
cascades, although he himself did
not use that term.
Predators help keep a balance
in a food web by regulating the
populations of other animals. When
ECOSYSTEMS 141
See also: Predator–prey equations 44–49 ■ The food chain 132–133 ■ The ecosystem 134–137 ■ Energy flow through
ecosystems 138–139 ■ Evolutionarily stable state 154–155 ■ Biodiversity and ecosystem function 156–157
different trophic levels. They he described changes in food

concluded that populations of webs that were brought about
producers, carnivores, and by the experimental removal
decomposers are limited by their of predatory starfish from the
respective resources. Competition intertidal zone in Washington
occurs between species on each of State. The concept of trophic
these three trophic levels. They also cascades is now generally accepted,
found that herbivore populations although debate continues as to
are seldom limited by the supply of how widespread they are.
plants, but are limited by predators,
so they are unlikely to compete Top-down cascades
with other herbivores for common This type of cascade is clearly
Ocher starfish prey on sea creatures resources. The paper highlighted demonstrated when a food chain
such as mussels and limpets. In a the important role of top-down is interrupted by the removal of
famous experiment, Robert Paine took forces (predation) in ecosystems, a top predator. The ecosystem
them out of their rock pools to observe and bottom-up forces (food supply). may continue to function despite
the effect on the rest of the food web.
American ecologist Robert the shift in species composition;
Paine was the first to use the term alternatively, the removal of one
they attack and eat prey, they affect “trophic cascade” when, in 1980, species may lead to the ❯❯
the number and behavior of that
prey—since prey move away when
predators are present. The impact Predators eat
of a predator can extend down to the
herbivore prey
next feeding level (trophic level),
affecting the population of the prey’s
own food source. In essence, by Bottom-up cascade Top-down cascade
controlling the population density
and behavior of their prey, predators Predators move into the If predators are
indirectly benefit and increase the area and numbers increase removed …
abundance of their prey’s prey.
Indirect interaction that occurs
across feeding levels is described
by ecologists as a trophic cascade.
By definition, trophic cascades must
Herbivore population
cross at least three feeding levels. Prey population increases
increases
Four- and five-level trophic cascades
are also known, although these are
less common.
Controlling factors
In 1960, the American ecologist Overgrazing brings
Increased rainfall
Nelson Hairston and his colleagues habitat change and loss
encourages vegetation
Frederick Smith and Lawrence of species richness
Slobodkin published a key paper
entitled “Community Structure,
Population Control, and Competition,”
which examined the factors that Herbivore prey eat plants
control populations of animals on
142 TROPHIC CASCADES

ecosystem’s breakdown. In the US, benthic invertebrate species. This
a trophic cascade on the coast of in turn led to an increase in floating
southern New England is believed nutrients, which ultimately boosted
to be responsible for the die-off of phytoplankton rather than species
saltmarsh habitat. Recreational on the sea floor. The net effect of
anglers have reduced the number the crabs’ arrival was to transfer Just as a deer herd lives
of predatory fish to such an extent nutrients from the sea floor to the in mortal fear of its wolves,
that the number of herbivorous water column—the water between so does a mountain live
crabs has expanded dramatically. the sediment and the surface—and in mortal fear of its deer.
The resulting increase in the to degrade the ecosystem. Aldo Leopold
consumption of marsh vegetation
has had a knock-on effect on other Bottom-up cascades
species that depend on it. If a plant—a primary producer—
Trophic cascades can also be is removed from an ecosystem, a
disturbed by the introduction and bottom-up cascade may result. For
spread of a nonnative species, as example, if a fungal disease causes
happened when the omnivorous grass to die-off, a rabbit population plants support more herbivores and
mud crab indigenous to waters that depends on it will crash. In more predators. This is in contrast to
on the east coast of North America turn, the predators that eat rabbits top-down cascades, in which more
and Mexico became common in will starve or be forced to move predators lead to fewer herbivorous
the Baltic Sea in the 1990s. Crabs, away, and the entire ecosystem prey and a greater mix of plants.
which are keystone species in could break down. Conversely,
many coastal food webs, feed on if planting or conservation efforts Beetles, ants, and moths
benthic (seafloor) communities— boost the mix of plant life, more Investigating trophic cascades
bivalves, gastropods, and other herbivores (including the pollinators in four-tier systems is more difficult,
small invertebrates—with that help plants reproduce and because predators at the top
devastating efficiency, creating spread) will be attracted, and with feeding level may eat predators at
a strong top-down cascade. The them more predators. the level below and also herbivores
increase in the number of mud In the bottom-up model, the below them, so the relationships
crabs in the Baltic, where there are responses of herbivores and their become very complex. In 1999,
no equivalent predators, resulted predators to increased plant variety researchers studying trophic
in a dramatic decline in the mix of follow in the same direction: more cascades in tropical rain forest in
Costa Rica got around this problem
by studying a system of three
trophic levels of invertebrates, in
which the top predator—a clerid
beetle—ate the predatory ants
in the level below it, but not the
herbivores in the level below that.
When the number of predatory
beetles in the study area was
increased, the population of
predatory ants fell dramatically.
This reduced the pressure on
dozens of species of herbivorous
Californian yellow bush lupines

are fast growing and invasive. The
plant can upset the ecosystem by
causing elevated nitrogen levels that
attract nonnative species.
ECOSYSTEMS 143
Early humans
and megafauna
In the last 60,000 years, which
includes the end of the last ice
age, about 51 genera of large
mammals became extinct in
North America. Most were
herbivores, including ground
sloths, mastodons, and large
armadillos, but many were
carnivores, such as American
lions and cheetahs, scimitar
cats, and short-faced bears.
Many of the extinctions
occurred between 11,500 and
10,000 years ago, shortly after
the arrival and spread of the
Steller’s sea cow was a giant cascade, triggered by the hunting Clovis people, who were
sirenian discovered by the naturalist to virtual extinction of sea otters for hunters. One of the most
Georg Steller in 1741. Its extinction is convincing theories about
the fur trade. The over-exploitation of
the cause of debate: was it hunted to these developments is the
death, or did its food source disappear? sea otters allowed the population “second-order predation
of sea urchins, their usual prey, to hypothesis,” which suggests
rise above a critical threshold. Sea that the humans triggered a
invertebrates, which therefore ate urchins eat kelp, so the growth of trophic cascade. The people
more vegetation. The leaf area their population led to a collapse in killed the large carnivores,
of the plants in the study was the extent of kelp forests—the sea which competed with them
consequently reduced by half. cows’ food source. Although the for prey. As a result, predator
Not all the “players” in trophic sea cows themselves were not numbers were reduced
cascades are obvious or visible. being hunted, they soon became and prey populations rose
Some are tiny and live underground. extinct. Understanding how such disproportionately, resulting
For example, yellow bush lupines— interventions, and the introduction of in overgrazing. The vegetation
plants that live on the Californian nonindigenous species, can damage could no longer support the
coast—are consumed by the trophic cascades is vital in shaping herbivores, with the result
caterpillars of ghost moths, which conservation measures today. ■ that many herbivores starved.
eat the lupines’ roots. In turn,
nematodes—wormlike
invertebrates—parasitize the
caterpillars. If these nematodes
are in the soil, they will limit the
population of caterpillars, and fewer
of the lupines’ roots will be affected. Herbivores are
usually expected to be
Extinction events well fed and carnivores
In extreme cases, a trophic cascade are usually expected
can lead to species extinction—as to be hungry.
in the case of Steller’s sea cows, Lawrence Slobodkin
marine mammals that once lived in Cave paintings in Altamira, Spain,
the Bering Strait but were declared show the importance of bison to
early humans. The wild population
extinct in 1768. It has recently been became extinct in 1927, but captive
argued that this extinction was herds have since been reintroduced.
caused by a calamitous trophic
ISLANDS ARE
ECOLOGICAL
SYSTEMS
ISLAND BIOGEOGRAPHY
146 ISLAND BIOGEOGRAPHY
IN CONTEXT
KEY FIGURES
Robert H. MacArthur
(1930–72), Edward O. Wilson
(1929–)
BEFORE
1948 Canadian lepidopterist
Eugene Munroe suggests a
correlation between island
size and butterfly diversity in
the Caribbean.
AFTER
1971–78 In the US, biologist
James H. Brown studies
mammal and bird species
variety on forest “islands” in
the Great Basin of California
and Utah.
2006 Canadian biologists
Attila Kalmar and David Currie
study bird populations on
346 oceanic islands and
discover that species variety
depends on climate as well as
area and isolation.
I
sland, or insular, biogeography Mangrove-fringed islands in the
examines the factors that Florida Keys—now protected for their
affect the species richness of diverse range of marine and terrestrial
life—were the focus of research to test
isolated natural communities. the island biogeography theory.
Charles Darwin, Alfred Russel
Unless we preserve Wallace, and other naturalists had
the rest of life, as a written about island flora and fauna pools within a dry landscape, or
sacred duty, we will be in the 19th century. Their studies fragments of mountain forest
endangering ourselves were conducted on actual islands between nonforested valleys.
by destroying the home in the ocean, but the same methods In the mid-20th century,
in which we evolved. can be used to look at any patch ecologists began more intensive
Edward O. Wilson of suitable habitat surrounded by studies into species distribution
unfavorable environment that on different islands, and how and
limits the dispersal of individuals. why they varied. In the US,
Examples include oases in the biologists Edward Wilson and
desert, cave systems, city parks in Robert MacArthur constructed the
an urban environment, freshwater first mathematical model of the
ECOSYSTEMS 147
See also: Evolution by natural selection 24–31 ■ Predator-prey equations 44–49
■ Field experiments 54–55 ■ The ecosystem 134–137
Random dispersal of organisms to islands
Mainland
Island 1
Robert H. MacArthur
Island 3
Island 2 Born in Toronto, Canada, in
An island’s size and distance from the mainland both 1930, and later relocating to
affect its species richness. Islands closer to the mainland Vermont in the US, Robert
will receive more random dispersion of organisms; the larger
MacArthur originally studied
island gets the most, the island furthest away gets the least.
mathematics. In 1957, he
received his Ph.D. from Yale
University for his thesis
factors at play in island ecosystems because if native species are exploring ecological niches
and, in 1967, they outlined a new pushed out of prime habitat by new occupied by warbler species
in conifer forests. MacArthur’s
theory of island biogeography. immigrants, they have a better
emphasis on the importance
Their theory proposed that each chance of finding an alternative,
of testing hypotheses helped
island reflected a balance between albeit imperfect (“suboptimal”) transform ecology from an
the rate of new species arriving habitat. Larger islands are also exclusively observational
there and the rate at which existing likely to have a greater variety of field to one that employed
species become extinct. For example, habitats or microhabitats in which experimental models as well.
a habitable but relatively empty to accommodate new immigrants. This methodology is reflected
island would have a low extinction A combination of variety and lower in The Theory of Island
rate since there are fewer species rates of extinction produces a Biogeography, which he
to become extinct. When more greater species mix than on a small coauthored with Edward O.
species arrive, competition for island—the “species-area effect.” Wilson. MacArthur received
limited resources increases. At a The actual species in the mix will awards throughout his career,
certain point, smaller populations change over time, as a result of and was elected to the
will be outcompeted, and the rate colonization and extinction, but will National Academy of Sciences
of species extinction will rise. An remain relatively diverse. in 1969. In 1972, he died of
renal cancer. The Ecological
equilibrium point occurs when the
Society of America awards a
species immigration rate and the Monitoring mangroves biennial prize in his name.
rate of those becoming extinct are In 1969, Wilson and his student
equal; this may remain constant Daniel Simberloff conducted a field Key works
until a change occurs in either rate. experiment that tested the theory
The theory also proposes that on six small mangrove islands in 1967 The Theory of Island
the rate of immigration depends the Florida Keys in the US. They Biogeography
on the distance from the mainland, recorded the species living there, 1971 Geographical Ecology:
or another island, and declines with then fumigated the mangroves to Patterns in the Distribution
increased distance. The area of an remove all the arthropods, such as of Species
island is a further factor. The larger insects, spiders, and crustaceans.
it is, the lower its rate of extinction, In each of the next two years, ❯❯
148 ISLAND BIOGEOGRAPHY

island of Cuba, because economic
sanctions since the 1960s have
The water acts as meant that fewer boats (and their
Islands are a physical barrier, lizard stowaways) dock in Cuba.
surrounded by water. preventing many species
from entering or leaving. “Island” habitats
In the early 1970s, American
biologist James H. Brown applied
the Wilson–MacArthur model to
“islands” of coniferous forest on 19
mountain ridges in the Great Basin
of California and Utah. The ridges
As species arrive are separated from each other by
Islands are and depart, island a vast sagebrush desert. Brown
ecological systems. populations evolve. discovered that the diversity and
distribution of small mammals
(excluding bats) in the isolated
forests could not be explained in
terms of an equilibrium between
they counted returning species floating vegetation). Most of these colonization and extinction. Some
to observe their recolonization. factors apply to any similar, isolated species had become extinct, but
The Florida Keys experiment habitats, not just actual islands. no new species had arrived for
showed that distance did indeed The impact of humans—who millions of years, so Brown dubbed
play an important role: the farther probably began visiting isolated the mammals “relicts.” A few years
an island was from the mainland, islands in the Pacific at least 3,000 later, his analysis of resident bird
the fewer invertebrates returned to years ago—has sometimes been populations on the ridges revealed
recolonize the area. dramatic. In recent centuries, that new bird species had arrived
New waves of immigration can, people took dogs, cats, goats, from larger, similar forests in the
however, save even faraway island and pigs with them when they Rocky Mountains to the east and
species from extinction. This is colonized islands in the Pacific in the Sierra Nevada to the west.
more likely to happen with certain and elsewhere; inadvertently, they Brown concluded that certain
bird species—which can travel long also carried rats on their boats. species groups—especially those
distances quickly—than with, for On many islands, rats ate the eggs that fly—are more likely to be
example, small mammals. There is of breeding seabirds and the successful immigrants than others.
also the so-called target effect, seeds of endemic plants, some of
where some islands are more which grew nowhere else. On the
favored destinations because of the Galapagos Islands, dogs ate
habitat they provide. Given the tortoise eggs, native iguanas, and
choice of a treeless island and one even penguins. Goats competed
with woodland, a tree-nesting bird with Galapagos tortoises for food
will naturally opt for trees. and wiped out up to five species of Destroying rainforest
plant on the Santiago Islands. for economic gain
Human impact The arrival of humans, however, is like burning a
The key factors influencing the has not always reduced species Renaissance painting
species mix on an oceanic island richness on islands. Researchers to cook a meal.
are its degree of isolation, how long discovered the important role of Edward O. Wilson
it has been isolated, its size, the ship-assisted colonization of
suitability of its habitat, its location islands in the Caribbean. Despite
relative to ocean currents, and its relatively small size, Trinidad,
chance arrivals (for example, for instance, has more species of
organisms washed up on mats of anole lizards than the much larger
ECOSYSTEMS 149
The rebirth of
Krakatau
In 1883, volcanic eruptions
devastated the Indonesian
island of Krakatau, wiping out
flora and fauna on the island
and nearby Sertung and
Panjang. By 1886, mosses,
algae, flowering plants, and
ferns had returned to Krakatau,
borne either on the wind or as
seeds on the surf. The first
young trees emerged in 1887;
various insect species, and a
single lizard, were discovered
in 1889. Recent research shows
that the level of immigration
to Krakatau and its neighbors
peaked during the period of
forest formation, from 1908 to
1921, but extinctions were at
their height when the dense
Ecologists also studied the diversity Central Park in Manhattan, New York
tree canopy prevented sunlight
of beetles and flies in nine parks of City, is an “island” in an urban setting.
Its checklist includes 134 bird species, from reaching the forest floor,
different sizes in Cincinnati, Ohio. between 1921–33. Although
197 insect, 9 mammal, 5 reptile, 59
Area was the best predictor of fungi, and 441 plant species. the immigration of land birds
species richness, but when the and reptiles has almost
ecologists coupled their findings stopped, new species of land
with data on population sizes, they activity, the island theory has also mollusk and many insect
calculated that an increased size of encouraged the creation of wildlife groups are still arriving from
parkland acts primarily to reduce corridors. These link areas of Sumatra and Java, both just
extinction rates rather than to suitable habitat, which helps under 28 miles (45 km) away.
provide habitats for new species. maintain ecological processes—for
example, allowing animal movement
Conservation practices and enabling viable populations to
Soon after the island biogeography survive—without requiring a great
theory was developed, ecologists expansion of protected areas. ■
began to apply it to conservation.
Nature reserves and national parks
were seen as “islands” in landscapes
altered by human activity. When
first creating protected areas,
ecologists debated the optimum
size: was one big reserve better than I will argue that every
several smaller ones? As the island scrap of biological
theory shows, biodiversity depends diversity is priceless …
on a number of factors, and different Edward O. Wilson
species benefit in different settings. Krakatau’s deadly eruption sent
A sizable mammal will not survive up an ash cloud 50 miles (80 km)
high that altered global weather
in a small reserve, but many small patterns and caused a temperature
organisms will thrive there. In drop of 2.2°F (1.2°C) for five years.
places under pressure from human
150
IT IS THE CONSTANCY
OF NUMBERS
THAT MATTERS
ECOLOGICAL RESILIENCE
T
he capacity for ecosystems population changes are not as
IN CONTEXT to recover following a important as whether the whole
disturbance—such as a system is being fundamentally
KEY FIGURE
large fire, flood, hurricane, severe altered. The resilience of a system
Crawford Stanley
pollution, deforestation, or the can be described either by the time
Holling (1930–)
introduction of an “exotic” new it takes to return to equilibrium
BEFORE species—is known as ecological after a big shock or by its capacity
1859 Charles Darwin resilience. Any of these impacts to absorb disturbance.
describes the interdependence can upset food webs, often One example that Holling studied
between species as an dramatically, and human activity was the fisheries of the Great Lakes
“entangled bank.” is responsible for an increasing in North America. A large tonnage
number of them. of sturgeon, herring, and other fish
1955 In the US, Robert was harvested in the early decades
MacArthur proposes Staying resilient of the 20th century, but overfishing
a measure of ecosystem Canadian ecologist Crawford dramatically reduced the catches.
stability that increases as Stanley Holling first proposed the Despite subsequent controls on
the number of interactions idea of ecological resilience to fishing, populations in the Great
between species multiplies. describe the persistence of natural Lakes did not recover. Holling
systems in the face of disruptive
1972 In contrast with changes. Holling argued that
MacArthur, Australian natural systems require stability
ecologist Robert May and resilience, but—contrary
argues that more diverse to what previous ecologists had
communities with more assumed—these are not always
complex relationships may be the same qualities. Ecosystems are dynamic—
less able to maintain a stable A stable system resists change constantly changing and
balance between species. in order to maintain the status quo, inherently uncertain, with
but resilience involves innovation potential multiple futures …
AFTER Crawford Stanley Holling
and adaptation. Holling wrote
2003 Australian ecologist that natural, undisturbed systems
Brian Walker works with are likely to be continually in a
Crawford Holling to refine transient state, with populations
the definition of resilience. of some species increasing and
others decreasing. However, these
ECOSYSTEMS 151
See also: The food chain 132–133 ■ The ecosystem 134–137 Energy flow
through ecosystems 138–139 ■ Trophic cascades 140–143
■
The role of budworm
Spruce budworm caterpillars
have devastated balsam fir
forests in eastern North
America six times since
the 18th century. Holling
described this process as
having two very different
states: one with young,
fast-growing trees and few
budworms; and one with
mature trees and very large
numbers of budworms.
Between outbreaks of
budworms, young balsam fir
grow alongside spruce and
white birch trees. Eventually,
the fir becomes dominant.
A combination of this
dominance and a sequence
of very dry years stimulates
a huge increase in the
budworm population. The
mature fir is destroyed, giving
the spruce and birch an
suggested that the intense fishing A thick green scum of algae covers opportunity to regenerate.
had progressively reduced the parts of Lonar Lake, in Maharashtra, By keeping the balsam fir
resilience of the ecosystem. India. Algae thrive in high-nutrient in check, the budworm also
conditions, but decomposing algae maintains the spruce and
Holling argued that ecological consume oxygen, and depleted levels of
resilience is not always positive. birch. Without it, the fir trees
oxygen lead to fewer fish surviving.
If a freshwater lake experiences would crowd out the others.
a large input of nutrients from So the system is unstable but
agricultural fertilizers, for example, components of the ecosystem at the same time resilient.
it will become eutrophic: algae will can change. One example is the
thrive, depleting the lake’s oxygen disappearance of most American
and making it unsuitable for fish. chestnuts from forests in eastern
Such a lake may be resilient, but it North America, which was largely
will become less biodiverse. Holling compensated for by the expansion
claimed that three critical factors of oaks and hickories. For Holling,
determine resilience: the most a this counted as resilience, because
system can be changed before although the exact mix of tree
crossing a threshold that makes species had changed, broad-leaved
total recovery impossible; the ease forest still remained.
or difficulty in making a big change Ecologists now understand that
to the system; and how close to the ecosystems can have more than
threshold a system is currently. one stable state. In Australia, for
example, woodlands dominated by Spruce budworm larvae in
Changing states mulga trees can exist in a grass- Quebec, Canada, feed voraciously
According to Holling’s view, rich environment that supports on fir and spruce before they
resilience at the ecosystem level is sheep-farming, or in a shrub- pupate. Moths emerge about a
enhanced by its populations not dominated environment that is month later, ready to mate.
being too rigid—meaning that the totally unsuitable for sheep. ■
152
POPULATIONS ARE
SUBJECTED TO
UNPREDICTABLE FORCES
THE NEUTRAL THEORY OF BIODIVERSITY
B
iodiversity is shaped
IN CONTEXT globally by new species
appearing and others
KEY FIGURES
becoming extinct. Community
Hal Caswell (1949–),
ecology has traditionally held that
Stephen P. Hubbell (1942–) Caswell made a bold attempt
interactions between species play
BEFORE a vital role in determining this to create a neutral theory of
1920 Frederic Clements process. If two species compete for community organization.
describes how plant species similar resources, for example, either Stephen P. Hubbell
are associated with each other the stronger pushes the weaker
in communities. to extinction, or each is driven into
a narrower niche of specialism.
1926 Henry Gleason proposes In 1976, however, American
that ecological communities ecologist Hal Caswell proposed
are organized more randomly. a “neutral” theory of biodiversity. It
maintained that ecologically similar Neutral theories of biodiversity have
1967 Richard Root introduces species are competitively equal, and dominated community ecology in
the concept of the ecological whether species become common recent years. However, an Australian
guild—a group of species or rare is down to chance processes. study of coral reefs, published in
exploiting resources in 2014, focusing on once-dominant
similar ways. The “null” model species that have been almost lost
AFTER In the early 2000s, American to overfishing, did not support the
ecologist Stephen P. Hubbell theory. According to Hubbell,
2018 A review headed
developed a mathematical model species are interchangeable, so
by Dutch ecologist Marten
known as the “null” hypothesis, others should have increased to take
Scheffer suggests that, published in The Unified Theory their place. The fact that this did not
although species that use of Biodiversity and Geography (2001), happen in this case suggests that
the same resources may that supported Caswell’s theory. the neutral theory is flawed. The
be competitively equivalent, He tested his model by studying question of what maintains diversity
they may also differ according real communities. remains an open one. ■
to their response to stress-
inducing factors, such as See also: Human activity and biodiversity 92–95 ■ Island biogeography 144–149
drought or disease. ■ Climax community 172–173 ■ Open community theory 174–175
ECOSYSTEMS 153
ONLY A COMMUNITY
OF RESEARCHERS HAS
A CHANCE OF REVEALING
THE COMPLEX
BIG ECOLOGY
WHOLE
A
n in-depth understanding affect hydrology, biodiversity, and
IN CONTEXT of ecosystems requires carbon dynamics—the way carbon
long-term study. In 1980, and nutrients move through the
KEY ORGANIZATION
the US National Science Foundation ecosystem. There are many other
National Science
set up six Long Term Ecological long-term research sites worldwide
Foundation (created 1950)
Research (LTER) sites to study with researchers logging data on
BEFORE long-term, large-scale ecological ecosystems. With free access to
1926 Russian geochemist phenomena. There are currently the information, the research can
and mineralogist Vladimir 28 sites, five of which have been be easily disseminated globally. ■
Vernadsky formulates the running since 1980. Ecologists are
theory of the biosphere amassing datasets that will enable
in which everything on in-depth knowledge to be shared.
Earth lives.
A forest ecosystem
1935 Pioneering British One of the six original research
ecologist Arthur Tansley sites is Andrews Forest in Oregon.
defines an ecosystem It provides a good example of a
as encompassing all the temperate rain forest, enjoying
interactions between mild, wet winters and cool, dry
a group of living creatures summers. With 40 percent being
and their environment. old-growth conifer forest, there is a
high degree of biodiversity across
AFTER its forest, stream, and meadow
1992 At the Earth Summit ecosystems. Ecologists have
in Rio de Janeiro, there is recorded thousands of species of
international consensus insects, 83 bird species, 19 conifer
on the importance of species, and 9 species of fish. Log decomposition is being studied
protecting the biosphere. Projects aim to observe how land- over a 200-year period at six old-growth
use (such as forestry) and natural forest sites in Andrews Forest, Oregon.
1997 The Kyoto Protocol phenomena (fires, floods, climate) The experiment began in 1985.
to reduce greenhouse gas
emissions is signed by See also: The ecosystem 134–137 ■ The biosphere 204–205 ■ Sustainable
192 countries. Biosphere Initiative 322–323 ■ Ecosystem services 328–329
154
THE BEST STRATEGY

DEPENDS ON WHAT
OTHERS ARE DOING
EVOLUTIONARILY STABLE STATE
T
he field of behavioral
IN CONTEXT ecology seeks to explain
how the behavior of
KEY FIGURE Animals come into animals—what they eat, how they
John Maynard Smith conflict with each other socialize, and so on—has evolved
(1920–2004) over food, territory, and to suit their particular environment.
mate selection. The driving force is natural selection
BEFORE
1944 Mathematician John von because the environment favors
Neumann and economist individuals with certain genes—
Oskar Morgenstern use a some genes are “better” for certain
theory of games of strategy to situations and not for others—
devise a mathematical theory which are then passed on to
of economic and social offspring. Because the behavior of
animals is influenced by genes,
organization.
behavior must be influenced by
They have evolved to
1964 British biologist W.D. natural selection as well.
react to the behavior of
Hamilton applies game theory other animals in certain
to the evolution of social preprogrammed ways. Adaptive behavior
behavior in animals. In 1972, British evolutionary
biologist John Maynard Smith
1965 Hamilton uses game introduced a theory known as the
theory to describe the evolutionarily stable strategy
ecological consequences (ESS), that helped explain how
of natural selection. behavioral strategies appear by
natural selection. Just as factors
1976 Richard Dawkins
such as food and temperature can
popularizes the idea of affect animals, so can the behavior
evolutionarily stable strategies. of other species. Maynard Smith
The best strategy
AFTER depends on what suggested that an ESS adapts to
1982 John Maynard Smith others are doing. the behavior of other animals, and
applies the theory to evolution, cannot be beaten by competing
sexual biology, and life cycles. strategies, thus giving animals the
best chance to pass on their genes.
He argued that only natural
ECOSYSTEMS 155
See also: Evolution by natural selection 24–31 ■ The selfish gene 38–39 ■ Predator–prey equations 44–49 ■ Ecological
niches 50–51 ■ Trophic cascades 140–143 ■ Biodiversity and ecosystem function 156–157
Behavior arising from conflicts behavior can be quantified, balance between two or more
over space and territory might emerge so biologists can work out which strategies within the system
as evolutionarily stable strategies. Fruit strategies are likely to be most as a whole. The overall balance
bats jostle for the best spots in the
trees, with alpha males driving weaker
stable by using mathematical is therefore better called an
bats down to lower branches. models (see box). If the model evolutionarily stable state, and
does not match the behavior not a strategy. Such a balance
of animals in the real world, then emerges when all individuals have
selection could upset this balance— it suggests that stability has equal fitness: they pass on their
hence why an ESS is “stable”—and not evolved. genes to the same extent. The
that these behaviour patterns are In real rather than hypothetical state remains stable, even when
genetically preprogrammed. ecosystems, it is not a single there are minor changes in the
ESS has its roots in game theory: strategy that is stable, but the animal’s environment. ■
a mathematical way of working out
the best strategy in a game. Many The hawk-dove “game”
examples of how animals behave
emerge as being evolutionarily The simplest demonstration in posturing. Which strategy
stable strategies, such as territorial of John Maynard Smith’s would be better for passing
behavior and hierarchies. For evolutionarily stable strategy on genes? Maynard Smith and
example, the genetically pre- (ESS) concerns a hypothetical his collaborators devised a
programmed “rules” of “if resident, response to aggression known mathematical model to provide
fight and defend” or “if visiting, give as the hawk-dove “game.” In the answer, and—in this
in and retreat,” which would help this, individuals can either be instance—being more hawkish
animals retain territory, combine hawkish and fight until badly than dovish emerged as the
to make territorial behavior an ESS. injured, or dovish and posture, ESS. It predicts a ratio of seven
but then retreat. Hawks will hawks for every five doves,
outmatch doves, but could be which is equivalent to any one
Balancing strategies seriously harmed in a fight with individual being hawkish
The payoff that an individual another hawk. Doves routinely seven-twelfths of the time, and
animal gains—or the price it risks escape injury, but waste time dovish five-twelfths of the time.
paying—by displaying a particular
156
SPECIES MAINTAIN
THE FUNCTIONING AND
STABILITY OF ECOSYSTEMS
BIODIVERSITY AND ECOSYSTEM FUNCTION
IN CONTEXT
KEY FIGURE
Michel Loreau (1954–)
BEFORE
1949 At the California Institute
of Technology in the US, the
first phytotron (research
greenhouse) is built to study
how an artificial ecosystem
can be manipulated.
1991 In the UK, an Ecotron,
a set of experimental
ecosystems in computer-
controlled units, is created at
Imperial College, London.
I
AFTER n an age when human A phytotron built in 1968 in North
2014 Leading ecologists in activities are rapidly eroding Carolina, US, now includes 60 growth
the US say that the effect of the complex mix of species in chambers, four greenhouses, and a
diversity loss on ecosystems controlled-environment facility for
different habitats, ecologists have studying plant diseases and insects.
is at least as great as—or even increasingly focused on how
greater than—that of fire, biodiversity loss affects the way
drought, or other drivers of ecosystems work. If species are Michel Loreau, director of the
environmental change. replaced or lost altogether, can an Centre for Biodiversity Theory
ecosystem remain intact—or does and Modeling in Moulis, France,
2015 A paper published in this damage ecosystem function? outlined diverse research; some
Nature provides evidence that Such questions were the focus looked more closely at species,
biodiversity increases an of the Biodiversity and Ecosystem others at what makes an ecosystem
ecosystem’s resilience in a Function (BEF) conference held in work. Loreau maintains that
broad range of climate events. Paris in 2000. More than 60 leading a new unified ecological theory
international ecologists, including is necessary to combat extreme
ECOSYSTEMS 157
See also: Mutualisms 56–59 ■ Keystone species 60–65 ■ The ecosystem
134–137 ■ Organisms and their environment 166 ■ Invasive species 270–273
When researching such issues,

ecologists have tended to use both
traditional observational fieldwork
and also sophisticated mathematical
models. More recently, research
Biodiversity loss… is likely has begun to incorporate the
to decrease the ability of manipulation of ecosystems
ecosystems to resist the in a more controlled way, on plots
effects of climate change. of land, for example, or within
Michel Loreau closed systems housed in giant
greenhouselike facilities called Habitat fragmentation
phytotrons. The experiments help
to establish what factors—such as Barro Colorado Island in the
numbers of species, or species type Panama Canal of Central
and dominance—affect ecosystems America was formed in 1914,
in the long term. Their findings when tropical rain forest was
environmental challenges. That, show that the effects of biodiversity flooded by damming, creating
he says, requires the integration on ecosystem functions are an isolated fragment of forest
of community ecology (the study of complex. While the most diverse surrounded by water. Since
how species interact in ecosystems) ecosystems tend to be the most 1946, the area has been
with ecosystem ecology (research productive, their success also studied in detail by biologists
into the physical, chemical, and depends on climate and soil fertility. of the Smithsonian Institution
and elsewhere to determine
biological processes that connect There is more to be learned
the effects of this habitat
organisms and their environment). about how plant diversity affects
fragmentation: species
soil processes, the role of microbe diversity on the island has
Complex cycles biodiversity in the soil, and the declined, and top predators are
Scientists of both disciplines firmly effects of mutualistic species, such among the most vulnerable
believe that biodiversity, especially as flowering plants and pollinating species. In the US, studies of
species and genetic diversity, is insects. Much has been achieved, habitat fragmentation and its
an important driver of ecosystem but questions remain, and the effects on diversity in the
functioning. Ecosystems are unifying theory that Loreau is Florida Keys led to Robert
powered by an input of energy and seeking has still to be devised. ■ MacArthur and E.O. Wilson’s
recycling of nutrients: plants and seminal Theory of Island
animals grow, die, and decompose, Biogeography (1967).
returning nutrients to the soil From such environments,
and restarting the cycle. These planners have learned
processes depend on the species important lessons about how
to conserve species in isolated
within the ecosystems, which in
One of the distinctive and patches of habitat—sometimes
turn depend upon one another as in the midst of cities—that are
they interact—as predators and fascinating features of
set aside as reserves. Barro
prey, for example. Many ecologists ecological systems is their Colorado, and places like it,
argue that a large variety of extraordinary complexity. have also provided vital
complementary species are needed Michel Loreau opportunties for study, where
to keep an ecosystem working and ecologists can explore how
make it resilient to change. Others changing species diversity
say that a few key species may be affects the functioning of an
more important to stop ecosystems ecosystem at every level.
from collapsing.
ORGANIS
IN ACHAN
ENVIRON
MS
GING
MENT
160 INTRODUCTION
Frederic Clements coins

Alexander von Humboldt The earliest forms Andreas Schimper shows the term “climax
sets out for Latin America, of ecological survey how the link between a community” for the
on an expedition that lays are conducted in plant’s physiology and process of stabilization
the groundwork for modern studies of wild fish external conditions over time in ecological
plant geography. by Stephen A. Forbes. is key to plant ecology. communities.
1799 1880S 1898 1916
1845 1895 1899
Pierre-François In showing how plants The changing vegetation

Verhulst formulates an are related to their of sand dunes inspires
equation to predict environments, Johannes Henry Chandler
population growth. Warming unites the fields of Cowles’ idea of
botany and ecology. primary succession.
T
he distribution of organisms represented the true birth of the While studying the vegetation
through space and time is field of ecology. Pioneers included growing on sand dunes along the
a fundamental interest of American naturalist Stephen A. shore of Lake Michigan in the
ecology. Early in the 19th century, Forbes, who studied wild fish 1890s, American botanist Henry
Prussian explorer Alexander von populations in the 1880s, and Chandler Cowles realized that there
Humboldt, a founding father of Danish botanist Johannes was a succession of plant species,
ecology, made detailed studies of Warming, who examined the with “pioneer” plants being
plant geography in Latin America. interaction between plants and replaced by others, which were in
Philip Sclater described the global their environment and introduced turn themselves supplanted. Fellow
distribution of bird species, and the idea of plant communities. American Frederic Clements used
Alfred Russel Wallace did the same The link between climate and the term “climax community”
for other vertebrates, proposing six a region’s dominant vegetation to describe the endpoint of this
zoogeographic regions that are type was set out by German succession. In 1916, he proposed
largely still in use today. botanist Andreas Schimper, who that global vegetation patterns
produced a worldwide classification could be thought of as “formations,”
Communities of vegetation zones in 1898. In the or large communities of plants—
Early fieldwork concentrated on early years of the 20th century, and the organisms that depended
the distribution and abundance ecologists devoted more attention on them—which reflected the
of organisms, but later in the 19th to the interrelatedness of all regional climate. In relatively wet,
century scientists increasingly organisms within an ecosystem, temperate regions, for example,
recognized that survey data could exemplified by Russian scientist deciduous forest may dominate,
also throw light on interactions Vladimir Vernadsky’s concept but grassland tends to dominate
between species. In a sense, this of the biosphere. in drier, more temperate areas.
ORGANISMS IN A CHANGING ENVIRONMENT 161
Robert May applies James H. Brown and Mathew Leibold’s

The concept of the chaos theory to Robert Maurer devise the “metacommunities”
“ecological guild” is predicting rates of concept of macroecology, concept looks at how
introduced in Richard growth and decline in in which ecological populations of a single
B. Root’s thesis on the animal population patterns are analysed species disperse
blue-grey gnatcatcher. dynamics. across large areas. and interact.
1967 1976 1989 2004
1957 1975 1988 1991
The first satellite goes Citizen science John Odling-Smee Ilkka Hanski outlines
into space, heralding enables Fred and Norah suggests that “niche his metapopulation
new technologies in Urquhart to discover constructors” actively theory for species in
wildlife tracking. where monarch change their environment. fragmented habitats.
butterflies go in winter.
Clements argued that these climax that “exploit the same class examples, from ancient oxygen-
communities were bound together of environmental resources,” producing cyanobacteria that altered
and could be thought of as single, regardless of how they do it. the composition of the atmosphere
complex organisms. in prehistoric times, to beavers
Clements was soon challenged New ideas creating wetlands.
by American botanist Henry Many new ideas enriched the
Gleason, who agreed that plant study of ecology in the late 20th Modern methods
communities could be mapped, but and early 21st centuries. The Traditionally, the task of monitoring
argued that since individual plant metapopulation concept was environmental change has been
species have no common purpose, advanced by the Finn Ilkka Hanski, the responsibility of academics
the idea of integrated communities who argued that a population of and professional ecologists, but
was invalid. His view found support a species is made up of differing, millions of interested amateurs now
in the 1950s, in the field studies of dynamic components. One part of provide enormous amounts of raw
Robert Whittaker and the numerical a population may become extinct, data on everything from flowering
research of John Curtis. while another thrives. The thriving dates to butterfly numbers, and
In 1967, American ecologist element may subsequently help from the state of coral reefs to the
Richard Root proposed the idea of reestablish the population that has breeding populations of birds. With
the “guild,” a group of organisms— died out. computer power to quickly process
closely related or otherwise—that In the process, British ecologist vast amounts of data, and with
exploit the same resources. Later, John Odling-Smee argued, so-called Earth’s ecology changing faster
ecologists James MacMahon and “niche-constructor” species create than ever, this “citizen science”
Charles Hawkins refined the a more favorable environment for looks set to become an invaluable
definition of a guild to species themselves—as seen in countless resource for ecology. ■
162
THE PHILOSOPHICAL STUDY

OF NATURE CONNECTS THE
PRESENT WITH THE PAST
THE DISTRIBUTION OF SPECIES
OVER SPACE AND TIME
IN CONTEXT
KEY FIGURE Species are Plants and animals move
Alexander von Humboldt distributed over time as Earth and its
(1769–1859) throughout the world. habitats change.
BEFORE
1750 Carl Linnaeus explains
that the distribution of plants
is determined by climate.
AFTER
1831–36 Charles Darwin The philosophical Scientists study
makes various observations study of nature where and how species
on the voyage of HMS Beagle, live now but also where
connects the they were before, and
confirming that many animals present to the past. what has changed.
living in one area are not found
in similar habitats elsewhere.
1874 British zoologist Philip
T
Sclater produces a description he distribution, or range, of species’ distributions, but the first
of the zoogeography (the biological communities and to make detailed studies of this
geographical distribution of species varies according to aspect of zoology was the Prussian
animals) of the world’s birds. many factors—including latitude, polymath Alexander von Humboldt,
climate, elevation, habitat, isolation, who traveled to Latin America with
1876 Alfred Russel Wallace
and the species’ characteristics. French botanist Aime Bonpland in
publishes his two-volume book The study of species distribution is 1799. Their five-year expedition
The Geographical Distribution called biogeography. Biogeography laid the basis of plant geography.
of Animals, which becomes is also concerned with how and Humboldt believed observation in
the definitive biogeography why the patterns of distribution situ to be paramount, and used
text for the next 80 years. change over time. sophisticated instruments to make
Early zoologists and botanists meticulous records of both plant
such as Carl Linnaeus were well and animal species, noting all the
aware of geographical variations in factors that could influence the

See also: Modern view of diversity 90–91 ■ Animal ecology 106–113 ■ Island
biogeography 144–149 ■ Big ecology 153 ■ Climate and vegetation 168–169
and New Zealand). The dividing line

between these last two regions,
which runs through Indonesia, is
still known as “Wallace’s Line.”
The unity of nature Plate tectonics

means the interrelationship Wallace also made some remarkable
of all physical sciences. discoveries from the fossil record.
Alexander von For example, he worked out that
Humboldt early rodents had evolved in the
Northern Hemisphere, moving via Alexander von
Eurasia into South America. Later,
Humboldt
in 1915, German geologist Alfred
Wegener proposed the radical idea Known as the “founder of plant
that the continents of South geography,” Humboldt also
America and Africa were once made valuable contributions
data. This holistic approach is best connected, which allowed the to geology, meteorology, and
illustrated in his highly detailed map spread of tapirs and other species. zoology. Born in Berlin in 1769,
and cross section of Chimborazo Wegener understood that the he started collecting plants,
mountain in Ecuador. distribution of species was in part a shells, and insects at an early
record of geological history. Species age. His expedition to Latin
Wallace’s contribution colonize new areas as conditions America in 1799–1804
Many 19th-century naturalists change, and over time have become encompassed Mexico, Cuba,
contributed to biogeographical separated by barriers such as Venezuela, Colombia, and
Ecuador, and his team broke
knowledge, but one of the most new oceans or mountain ranges.
the world altitude record when
significant was British naturalist Today, as human-made changes to they climbed to 19,285ft
Alfred Russel Wallace. After reading climate and the environment gather (5,878m) on Chimborazo.
Philip Sclater’s account of the global pace—creating new barriers—this Humboldt also speculated
distribution of bird species, Wallace understanding has taken on a new that volcanoes result from
set out to do the same for other and vital importance. ■ deep subterranean fissures,
animals. He examined all the factors investigated the decrease in
known at the time to be relevant, temperature with altitude, and
including changes in land bridges discovered that the strength
and the effects of glaciations. He of Earth’s magnetic field
produced maps to demonstrate decreases away from the
how vegetation influenced animal poles. The 23-volume work
ranges, and he summarized the detailing his expedition set a
distribution of all known families new standard for scientific
writing, cementing his fame.
of vertebrates.
Wallace then proposed six
Key works
zoogeographic regions, which are
still largely in use today: the Nearctic 1807 Essay on the Geography
(North America), Neotropics (South of Plants
America), Palearctic (Europe, north 1805–1829 Personal Narrative
Tapirs evolved in North America at
Africa, and most of Asia), Afrotropics least 50 million years ago. They spread of Travels to the Equinoctial
(south of the Sahara), Indomalaya to and now live in Central and South Regions of the New Continent
(South and Southeast Asia), and America, as well as southeast Asia, but During Years 1799–1804
Australasia (Australia, New Guinea, died out in North America.
164
THE VIRTUAL INCREASE

OF THE POPULATION IS
LIMITED BY THE FERTILITY
OF THE COUNTRY
THE VERHULST EQUATION
P
ierre-François Verhulst was did not take into account a larger
IN CONTEXT a Belgian mathematician population’s difficulty in finding
who, after reading Thomas food. He argued instead that “the
KEY FIGURES
Malthus’s An Essay on the Principle population gets closer and closer
Thomas Malthus
of Population, became fascinated by to a steady state,” in which the rate
(1766–1834), Pierre-François
human population growth. In 1845, of reproduction is proportionate to
Verhulst (1804–49) he published his own model for both the existing population and
BEFORE population dynamics, which was the amount of available food. In
1798 Thomas Malthus argues later named the Verhulst equation. Verhulst’s model, after the point of
that populations increase Although influenced by the maximum population growth—the
exponentially, based on a ideas of Malthus, Verhulst realized “point of inflection”—the growth
common ratio, whereas food that there was a major flaw in his rate becomes progressively slower,
supplies grow more slowly at predictions. Malthus had claimed gradually leveling off to reach the
a constant rate, leading to that human population tends to “carrying capacity” of an area—the
potential food shortages. increase geometrically, doubling number of individuals it can sustain.
at regular time intervals. Verhulst When visualized, Verhulst’s model
1835 Belgian statistician thought this to be too simplistic, produces an S-shaped curve, which
Adolphe Quetelet suggests reasoning that the Malthus model was later called a logistic curve.
that population growth tends
to slow down as population Practical demonstrations
density increases. Verhulst’s model was ignored for
several decades, partly because he
AFTER himself was not entirely convinced.
1911 Anderson McKendrick, However, in 1911, Scottish army
working as an army physician, The hypothesis of physician and epidemiologist
applies the Verhulst equation geometric progression Anderson McKendrick used the
to bacteria populations. can hold only in very logistic equation to forecast
special circumstances. growth in populations of bacteria.
1920 American biologist Pierre-François Verhulst Then, in 1920, Verhulst’s equation
Raymond Pearl proposes the was adopted and promoted in
Verhulst equation as a “law” America by Raymond Pearl.
of population growth. Pearl conducted experiments
with fruit flies and hens. He gave
a constant quantity of food to fruit

See also: Distribution of species over space and time 162–163 ■ Metapopulations
186–187 ■ Metacommunities 190–193 ■ Overpopulation 250–251
laid fewer eggs and, as their

fertility rate declined, the rate of
population growth slowly stabilized.
Variable strategies
Biologists are at the The two key variables in Verhulst’s
present time in no equation are the maximum capacity
way likely to suffer of a species to reproduce (r), and the
ostracism if they carrying capacity of the area (K).
venture to study Organisms are either r-strategists
human problems. or K-strategists. R-strategists, such Thomas Malthus
Raymond Pearl as bacteria, mice, and small birds,
reproduce rapidly, mature early, Malthus was born in Surrey,
and have a relatively short life. UK, in 1766, the seventh child
K-strategists, such as humans, of a prosperous family. After
elephants, and giant redwood studying languages and
trees, have a slower reproduction mathematics at the University
rate, take longer to mature, and of Cambridge, he took a post
flies kept in a bottle. Initially, their tend to live longer. Ecologists as curate of a rural church. In
fertility rate increased. However, study r-strategists, which are often 1798, he published an essay
as the population density grew, found in unstable environments, arguing that the rate of
competition for resources increased, to assess risks to their necessary increase in human populations
outstrips much steadier rises
and eventually reached a bottleneck. high reproduction levels, and study
in food production, leading to
After this, the flies’ fertility rate K-strategists in more predictable
inevitable starvation. Malthus
dropped; their numbers continued environments to ensure long-term went on to publish six further
to increase but slowly, and generally species survival. ■ editions of the essay, and he
the population level stabilized. made a number of visits to
Similarly, Pearl found that Europe to gather population
Fruit flies are small, common flies that
when the number of hens in a pen are attracted to ripe fruit and vegetables. data. In 1805, he was appointed
increased, the birds struggled to They are popular for laboratory studies Professor of History and
find enough food. As the space because they reproduce so quickly and Political Economy at the East
between them reduced, the hens are easy to cultivate. India Company College in
Hertfordshire. He became
increasingly involved in
debate about economic policy,
and criticized the Poor Laws
for causing inflation and
failing to improve life for the
poor. Malthus died in 1834.
Key works
1798 An Essay on the Principle

of Population
1820 Principles of Political
Economy
1827 Definitions in Political
Economy
166
THE FIRST REQUISITE

IS A THOROUGH
KNOWLEDGE OF
THE NATURAL ORDER
ORGANISMS AND THEIR ENVIRONMENT
T
he notion of a naturalist— analysis and experiments. These
IN CONTEXT someone who studies rounded ecological surveys created
organisms in the natural a picture of the natural order within
KEY FIGURE
world—dates back to ancient an environment. By shedding light
Stephen A. Forbes
Greece. Aristotle made copious on the interrelated effects of its
(1844–1930)
observations of wildlife, and his plant and animal life, they could
BEFORE work laid the foundations for later also help explain the distribution of
1799–1804 Alexander von naturalists. It was not until the species and variations over time. ■
Humboldt pioneers the field 19th century, however, that the
of biogeography in his travels potential of such surveys was
in Latin America. really understood.
1866 German naturalist The new study of ecology

Ernst Haeckel coins the term As naturalists undertook longer
“ecology” to describe the study field trips, the global distribution
of organisms in relation to of species became more apparent,
their environments. and the concept of ecology as a
science gained traction.
1876 After traveling One of the first scientists to
extensively, British naturalist employ ecological methods was
Alfred Russel Wallace American biologist Stephen A.
publishes The Geographical Forbes. In the 1880s, while
Distribution of Animals. studying fish in a Wisconsin lake,
he realized that survey data could
AFTER
be interpreted to give a picture of
1890s Frederic Clements interactions between different
proposes the notion of species—not just their abundance.
Satellite images enable ecologists to
ecological communities. observe large-scale changes easily. The
Forbes extended the scope of the green areas in this image of the Caspian
1895 In Ecology of Plants conventional survey, combining Sea are evidence of algal growth—the
Johannes Warming describes practical fieldwork with theoretical product of nutrient enrichment.
the impact of the environment
on the distribution of plants. See also: Classification of living things 82–83 ■ Animal ecology 106–113
■ Biodiversity and ecosystem function 156–157
PLANTS LIVE
ON A DIFFERENT
TIMESCALE
THE FOUNDATIONS OF PLANT ECOLOGY
P
lant ecology examines how
IN CONTEXT plants interact with one
another and with their
KEY FIGURE
environments. Danish botanist
Johannes Eugenius
Johannes Eugenius Warming first
Warming (1841–1924)
brought the sciences of botany and That land is a community
BEFORE ecology together in his book The is the basic concept
1859 Charles Darwin’s Ecology of Plants in 1895. He of ecology.
detailed descriptions of plants described how plants react to their Aldo Leopold
and animals in their natural surroundings, and how their life American ecologist
environment mark the start of cycles and structures relate to where
an appreciation of what is later they grow. The book introduced the
termed “ecology.” concept of plant communities, and
outlined how a group of species
AFTER interact and develop in reponse to
1935 British botanist Arthur the same local conditions.
Tansley publishes an article in biosphere, the parts of its surface
Ecology in which he defines Plants and ecosystems and atmosphere where all living
the term “ecosystem.” For many years, plant ecology organisms exist and interact.
and animal ecology were studied Plants are sensitive barometers
1938 American botanists separately, but in the early 20th of change within an environment.
John Weaver and Frederic century a more connected The study of their anatomy,
Clements further develop the perspective emerged. Important physiology, distribution, and
concepts of plant communities theories on plant communities and abundance, as well as their
and succession. succession—the process by which interactions with other organisms
an ecological community changes and their response to environmental
1995 David Attenborough’s over time—were established during factors, such as soil conditions,
television documentary “The this time period. In 1926, Russian hydrology, and pollution, can
Private Life of Plants” depicts geochemist Vladimir Vernadsky provide invaluable information
plants as dynamic influencers introduced the idea of Earth’s about the entire ecosystem. ■
of their environment.
See also: Climate and vegetation 168–169 ■ Ecological succession 170–171
■ The biosphere 204–205 ■ Endangered habitats 236–239 ■ Deforestation 254–259
168
THE CAUSES OF
DIFFERENCES
AMONG PLANTS
CLIMATE AND VEGETATION
T
hat different plants grow
IN CONTEXT in different climates was
likely common knowledge
KEY FIGURE
for as long as agriculture has
Andreas Schimper
existed; many cultures have
(1856–1901)
traded plants for thousands of years.
BEFORE However, the clear link between a
1737 Carl Linnaeus’s Flora region’s dominant vegetation type
Lapponica includes details of and climate was not categorically
the geographical distribution spelled out until German botanist
of Lapland plants. Andreas Schimper published his
ideas on plant geography in 1898.
1807 Alexander von Humboldt Botanists such as Carl Linnaeus
publishes his seminal Essay and Alexander von Humboldt had
on the Geography of Plants. written about plant distributions in
the 18th and early 19th centuries. “Flowering stones” (Lithops) are
AFTER The widely traveled Humboldt native to southern Africa, their thick,
1916 In Plant Succession: understood that climate was one fleshy leaves well suited to dry, rocky
an Analysis of the conditions. Related species also occur
of the key factors governing where in similar arid habitats in the US.
Development of Vegetation, plants did and did not grow.
Frederic Clements describes Schimper went one step further
how communities of species than Humboldt by explaining plant physiology (the functioning of
are indicators of the climate that similar vegetation types arise plants), it became the foundation of
in which they have matured. under similar climatic conditions the study of plant ecology. Schimper
in different parts of the world. explained that the connection
1968 “The Role of Climate
He then produced a global between the structures of plants
in the Distribution of classification of vegetation zones and the external conditions they
Vegetation,” by American that reflected this observation. faced in different places was the
geographers John Mather Schimper’s 1898 book Plant- key to what he described as
and Gary Yoshioka, explains geography upon a Physiological “ecological plant-geography.”
how temperature and rainfall Basis ran to 870 pages and is one Vegetation was divided into broad
alone are not enough to define of the largest ecology monographs tropical, temperate, arctic,
plant distributions. written by a single author. A mountain, and aquatic zones, then
synthesis of plant geography and subdivided further, according to

See also: Evolution by natural selection 24–31 ■ Ecophysiology 72–73 ■ The ecosystem 134–137 ■ The foundations of plant
ecology 167 ■ Biogeography 200–201 ■ Biomes 206–209
contrivances for regulating the were linked by warm temperatures

passage of water.” To illustrate this, and year-round moisture—traits of
he chose a type of vegetation with what he called a tropical rain forest.
tough leaves, short internodes (the Although the broad geographic
distances between the leaves along divisions devised by Schimper still
… the time is not far a stem), and leaf orientation parallel hold true, there is now a better
distant when all species or oblique to direct sunlight. This understanding of how vegetation
of plants and their type grew in various parts of the develops in response to many
geographical distribution world, where arid conditions meant different stimuli beyond simple
will be well known. that water was scarce. The name climatic differences. For example,
Andreas Schimper Schimper gave to these plants— measures of potential water
sclerophyll, from the Greek words evaporation into the atmosphere,
skleros (“hard”) and phullon (“leaf”)— water surplus, and water deficit,
is still used today. which can be combined in a
Epiphytes, plants that grow on moisture index, are more useful
the surface of other plants and determinants of plant distribution
derive their moisture and nutrients than simple temperature and
the prevailing climate. For example, from the air or rain, also fascinated rainfall figures. ■
tropical vegetation was divided into Schimper. He observed epiphytes
savanna, thorn-forest, woodland, such as Spanish moss growing in
Like other epiphytes, Spanish moss
tropical rain forest, or woodland the southern US and the Caribbean lives on other species but draws water
with a pronounced dry season, islands and similar species in and nutrients from the air rather than
according to whether the climate South America, South Asia, and from its host. It thrives in tropical and
was wet all year round, seasonally southeast Asia. He found that they subtropical environments.
wet, or mostly dry.
Adaptations for extremes

Schimper made a close study of
plant physiology—the structures of
plants and how they had adapted to
varying temperature and moisture
conditions. He was particularly
interested in plants growing in
extreme climatic conditions. Salty
environments, for example, require
plants to survive high levels of
soil and water salinity. Schimper
found that vegetation growing on
the coastal mangroves of Brazil, on
Caribbean and Sri Lankan beaches,
and in sulfur-emitting volcanic
craters in Java, were similarly
tolerant to salt.
Schimper also studied how
plants coped in the challenging
conditions of arid environments.
He found that plants growing in
hot, dry places had evolved “varied
170
I HAVE GREAT
FAITH IN A SEED
ECOLOGICAL SUCCESSION
IN CONTEXT
KEY FIGURE
Henry Chandler Cowles
(1869–1939)
BEFORE
1825 Adolphe Dureau de
la Malle coins the term
“succession” when describing
new growth in forest cuttings.
1863 Austrian botanist
Anton Kerner publishes a
study of plant succession in
the Danube river basin.
AFTER
1916 Frederic Clements
T
suggests that communities he Indiana Dunes comprise 15,000 years ago, there would only
settle into a climax, or stable a windswept section of have been bare sand around Lake
equilibrium, at the end of a shifting sand along the Michigan’s shore. Vegetation developed
succession period. in a physical gradient, with sand nearest
southern shore of Lake Michigan, the water and forests farthest back.
US. In 1896, American botanist
1977 Ecologists Joe Connell
Henry Chandler Cowles saw these
and Ralph Slatyer argue that
dunes for the first time, and so decomposing matter created
succession occurs in diverse began his career in the emerging favorable conditions for other plants.
ways, highlighting facilitation field of ecology. Dunes are among As these new plants died, even more
(preparing the way for later some of the planet’s least stable plants could grow.
species), tolerance (of lower landforms, and therefore changes Based on his observations,
resources), and inhibition to their ecology happen relatively Cowles developed the idea of
(resisting competitors). quickly. As Cowles walked among ecological succession, although
the dunes, he noticed that when groundwork for the concept had
certain plants died off, their been laid by earlier naturalists. In

See also: Field experiments 54–55 ■ The ecosystem 134–137 ■ Climax community 172–173 ■ Open community theory
174–175 ■ Biomes 206–209 ■ Romanticism, conservation, and ecology 298
Primary succession
The process of primary succession begins in barren
environments such as bare rock. Hardy species, usually lichens,
appear first and then give way to a stable climax community of
more complex and diverse life forms over hundreds of years.
Soil
Grasses,
Small annual shrubs,
plants and Grasses and and shade- Shade-
Bare rock Lichens lichens perennials intolerant trees tolerant trees
Hundreds of years
Pioneer species Intermediate Climax

species community
an 1860 address to members of the include pioneer plants (often lichens resulting from the pioneer species,
Middlesex Agricultural Society, and mosses), followed by grassy shrubs and oak, pine, and hickory
Massachusetts, Henry David plants, small shrubs, and trees. trees will begin to grow. As the
Thoreau had stated: “Though I do trees grow higher, shading out more
not believe that a plant will spring Life after disturbance of the underbrush, the grasses are
up where no seed has been, I have Secondary succession occurs after a replaced by plants able to survive
great faith in a seed.” disturbance that destroys plant life, with low sunlight, and, after around
such as a flood or a fire. The plant 150 years, the forest once more
Growth of an ecoystem life reestablishes itself and develops resembles the prefire community. ■
French geographer Adolphe Dureau into an ecosystem similar to the one
de la Malle is regarded as the first that existed before the disturbance.
person to use the term “succession” The stages of secondary succession
with reference to ecology when he are similar to those of primary
witnessed the progression of plant succession, although the ecosystem
communities after all the trees may start at different points in the I … found indisputable
were removed from a forest. Cowles process, depending on the level of evidence (a) that forests
provided a more formal articulation damage caused by the trigger. succeeded prairie, and
of his ecological succession theory, A common example of secondary (b) that prairie had
in The Ecological Relations of the succession occurs after a wildfire in
succeeded forest.
Vegetation on the Sand Dunes of oak and hickory forests. Nutrients
Lake Michigan, published in 1899. from burned plants and animals
Henry Allan Gleason
American ecologist
In this seminal paper, he proposed provide the right conditions for
the idea of primary succession— growth of annual plants. Pioneer
the gradual growth of an ecosystem grasses soon follow. After several
originally largely devoid of plant life. years, due at least in part to the
The stages of primary succession environmental and soil changes
172
THE COMMUNITY
ARISES, GROWS,
MATURES,
CLIMAX COMMUNITY
AND DIES
T
he term “climax community”
IN CONTEXT In every region, plants was first proposed in 1916
grow and develop by American botanist
KEY FIGURE
through a series Frederic Clements. He used it to
Frederic Clements
of successions. describe an enduring ecological
(1874–1945)
community that has reached a
BEFORE steady state, such as a naturally
1872 German botanist stable forest of old-growth trees
August Grisebach classifies that has not undergone or been
the world’s vegetation patterns subjected to any unnatural
in relation to climate. At each stage, they changes, such as logging.
become bigger,
1874 British philosopher more complex, and Regional communities
Herbert Spencer suggests that interconnected. In the 19th century, German
the human population can be botanists August Grisebach and
thought of as a giant organism. Oscar Drude were among those
who recognized that patterns of
1899 In the US, Henry Cowles vegetation around the world reflect
proposes that vegetation factors such as climate variations.
develops in stages, a process It was clear, for example, that the
called succession. Eventually the vegetation
typical vegetation in a wet, tropical
takes on the most
AFTER complicated interconnected climate was very different to that in
form the climate will allow. a dry, temperate climate. Then in a
1926 US ecologist Henry
landmark paper in 1899, American
Gleason argues that a climax
botanist Henry Cowles described
community is a coincidental
how plants colonized sand dunes
collection of individuals. around Lake Michigan in stages—
1939 British botanist Arthur or “successions”—of increasing
Tansley suggests there is not size and complexity.
Once a community In an influential book, Plant
a single climax community but reaches this
“polyclimaxes” responding to Succession (1916), Frederic
“climax,” vegetation Clements developed Cowles’s
various factors. stops changing. idea, which he combined with the
biogeographic thinking of the two

See also: The ecosystem 134–137 ■ The distribution of species over space and time 162–163 ■ Ecological succession
170–171 ■ Open community theory 174–175 ■ The ecological guild 176–177 ■ Biomes 206–209
The Sonora Desert is often seen as

an example of a climax community. It
has both winter and summer rains, so
its unique plants, which include the
tall saguaro cactus, are unusually lush.
German botanists to produce

a theory of the development of
natural communities.
Clements suggested that the
way to understand patterns of
vegetation across the world is
to think in terms of “formations.”
A formation is a large, natural
community of plants dominated
by a range of life forms that reflects
the regional climate. In each region,
plants go through stages or
successions until they reach the
most complex, highly developed
form of vegetation possible. Once
it finally reaches this climax, the
community stabilizes, in what
was later termed a “steady state,” stages of growth, he argued that for identifying such a community,
and stops changing. it can be considered as a single offered by American botanist Frank
Clements then proposed that complex organism. A community Egler in the 1950s, was never
climax communities are bound grows toward a climax in the claimed. Despite the difficulties,
together. Although an ecological same way that an individual ecologists continued to use the
community is made up of a develops through life stages. theory of a climax community to
multitude of plants at different Clements expanded the idea to decide how to respond to invasive
embrace all organisms in a “biome” species that threatened to disturb
that comprised “all the species of an established native community,
plants and animals at home in a and in recent decades Clements’s
particular habitat.” From this, ideas have regained support.
the idea of the ecosystem as a Succession remains a core
“superorganism” later developed. principle of ecology. In general,
For Clements, climates are early succession phases consist of
like genomes, and vegetation A fluctuating process fast-growing and well-dispersed
is like an organism whose Clements’s ideas were challenged species that are replaced by more
characteristics its from the start, although the idea of competitive species. Initially,
genome determines. a “steady state” proved influential ecologists thought that ecological
Christopher Elliott and dominated thinking about succession ended in what they
Philosopher of science ecosystems up until the 1960s. described as the climax phase,
However, scientists realized that when the ecosystem reached a
communities change constantly in stable equilibrium. However, it
response to conditions, and it is is now accepted that ecological
almost impossible to observe a true succession is a dynamic process
climax community. A $10,000 prize that is constantly in flux. ■
174
AN ASSOCIATION
IS NOT AN ORGANISM
BUT A COINCIDENCE
OPEN COMMUNITY THEORY
IN CONTEXT There is no evidence of

Plants grow according to
KEY FIGURE integrated development
their individual needs
between plants
Henry Allan Gleason
(1882–1975)
BEFORE
1793 Alexander von Humboldt
uses the word “association” to An ecological They grow randomly,
sum up the range of plant community is not influenced only by the
types in a particular habitat. an organism environmental conditions
1899 In the US, Henry Cowles
states that vegetation develops
W
in stages, in a process he calls hen American plant could see none of the integration
plant succession. ecologist Frederic proposed by Clements. Instead,
Clements proposed the Gleason believed that groups of
1916 Frederic Clements posits idea of climax communities in 1916, plants were random growths
the idea of a climax community he envisioned the community as a of individuals and species,
as a single organism. superorganism in which all plants responding to local conditions.
AFTER and animals interact to develop the
1935 Arthur Tansley coins community. A year later, American Individual needs
plant ecologist Henry Gleason Gleason maintained that the
the term “ecosystem.”
dismissed the idea; he argued that changes that occur during plant
1947 Robert H. Whittaker plant species have no common succession, as the composition
begins field studies that will purpose but merely pursue their of a community evolves, are
refute Clements’s holistic idea own individual needs. Gleason’s not integrated stages, as in the
of plant communities. hypothesis became known as the development of a single organism.
“open community” theory. The Rather, they are a combination of
1959 John Curtis boosts dispute initiated a debate that still responses from individual species
Henry Gleason’s reputation rages in ecological circles today. as they seek to meet their own
with numerical studies of Gleason did not deny that plant needs within a locality. “Every
prairie plant communities. communities could be mapped and species of plant,” Gleason argued,
their interactions identified, but he “is a law unto itself.” Gleason also

See also: The ecosystem 134–137 ■ The distribution of species over space and time 162–163 ■ Ecological succession
170–171 ■ Climax community theory 172–173 ■ The ecological guild 176–177 ■ Biomes 206–209
Diseases such as American chestnut

blight challenge the idea of a fully
integrated climax community, as the
loss of the dominant tree species should
cause the entire ecosystem to collapse.
during the 1930s as holism became

progressively supported by the idea
of the interactive “ecosystem.”
Nonetheless, as ecologists
continued to study the world, they
found more and more flaws in
Clements’s theory. In the 1950s, the
work of American plant ecologists
Robert H. Whittaker and John Curtis
showed how impossible it was to
identify communities as neat units
of holistic theory, and that the real
world was more nuanced and
complex. When it comes to studying
ecosystems in the field, Gleason’s
ideas seem to provide a better fit.
denied that there is any endpoint or need not worry too much about In the ensuing decades, while
climax community; he believed that disturbing the balance of the environmentalists continue to
communities are always changing. natural environment—because champion holistic ideas, ecologists
there is no balance. Gleason’s ideas have also increasingly incorporated
Changing opinions were therefore forgotten in the Gleason’s concepts into their work.
Gleason’s argument with Clements enthusiasm for developing ecology He is now considered to be one
caused quite a stir at the time. as a science. He became so of the most significant figures in
Clements seemed to be creating an frustrated that he gave up ecology 20th-century ecology. ■
overview in which natural patterns
of vegetation were determined by Henry Allan Gleason
clear rules, just as in Newtonian
science the movement of the planets Born in 1882, Henry Gleason classification. With botanist
is dictated by incontrovertible laws. studied biology at the University Arthur Cronquist, he co-wrote a
Clements and his supporters were of Illinois. He held faculty posts definitive guide to the plants of
able to look at the bigger picture, and conducted acclaimed early the northeastern US. He retired
while Gleason was viewed as a ecological research in Sand in 1950 but continued to write
reductionist, myopically intent on Ridge State Forest, Illinois. In and study. He died in 1975.
the details and challenging the the 1920s, Gleason’s theory of
entire idea of ecology as a science individualistic—rather than
controlled by laws. holistic—plant communities was Key works
not accepted by ecologists. This
Gleason appeared to be saying
rejection led Gleason to abandon 1922 “On the Relation between
that there are no patterns in nature:
ecology in the 1930s. He had Species and Area”
it is all random. Worse still, he was long held posts at the New York 1926 “The Individualistic
accused by some of justifying Botanical Garden and became Concept of the Plant
exploitative farming, since his famed for his work on plant Association”
ideas seemed to imply that man
176
A GROUP OF SPECIES
THAT EXPLOIT
THEIR ENVIRONMENT
IN A SIMILAR
THE ECOLOGICAL GUILD
WAY
E
cologists have long sought into its dry, scrubby chaparral
IN CONTEXT to understand how species environment. The thrasher’s
in a community interact to “niche” describes the aspects
KEY FIGURE
exploit resources. A key concept of its habitat for which it is
Richard B. Root (1936–2013)
in the explanation of this interplay suitably adapted.
BEFORE is the idea of guilds, first developed Root observed that the Blue-
1793 Alexander von Humboldt by American biologist and ecologist gray Gnatcatcher feeds on insects
uses the word “association” to Richard B. Root in 1967. that live on oak leaves. By
describe the mix of plant types Root had researched the way analyzing stomach contents, he
within a particular habitat. the Blue-gray Gnatcatcher exploits showed that several other birds
its ecological niche for his doctoral
1917 In the US, Joseph thesis. The concept of ecological
The Blue-gray Gnatcatcher is a
Grinnell coins the term “niche” niches dates back to 1917, when member of a guild of small birds that
to describe how a species fits American biologist Joseph Grinnell eat insects living on oak trees. Other
into its environment. used the term to describe how members of the guild include Hutton’s
the California Thrasher fitted Vireo and Oak Titmouse.
1935 British botanist
Arthur Tansley identifies
ecoystems—integrated biotic
communities—as fundamental
units of ecology.
AFTER
1989 In the US, James
MacMahon suggests that it
does not matter how ecological
guild members use resources.
2001 Argentinian ecologists
Sandra Diaz and Marcelo
Cabido propose grouping
species that have a similar
effect on their environment.

See also: Evolution by natural selection 24–31 ■ Predator–prey equations 44–49 ■ Optimal foraging theory 66–67 ■ Animal
ecology 106–113 ■ Open community theory 174–175 ■ Niche construction 188–189 ■ Metacommunities 190–193
also consume oak-leaf insects

and proposed that these oak-leaf
feeding birds could be grouped into Species
a “guild”—the “oak-foliage gleaners Different These species that
guild”—because they exploited the species may are linked share a
same resource. exploit by their resource
the same shared can be
resource. resource. grouped
Shared resources
Root defined a guild as a group of in a guild.
species that “exploit the same class
of environmental resources in a
similar way.” It does not matter
whether species in a guild are The guild concept was a major
related or not—all that matters is breakthrough in thinking about Richard B. Root
how they use their environment. connections between organisms
They do not even have to occupy in ecosystems. The theory implied American biologist and
ecologist Richard Root was
the same niche; they just have to that the entire functioning of an
born in Dearborn, Michigan,
use the same resource. ecosystem could be understood by in 1936. He grew up on a farm,
Guilds are typically identified identifying all the guilds within it. exploring nature and longing
by the food resource they have in Although that was potentially a to know “how the woods
common, although it could be any huge undertaking, ecologists have worked.” By the time he
other resource that they share. now managed to identify many completed his doctorate at the
Sharing the same resource means more guilds that confirm links University of Michigan, Root
that guild members often compete between species. For example, was already a knowledgeable
with one another, but they are not the birds of North America can ecologist. His 1967 thesis on
necessarily in constant competition. be grouped into guilds of gleaners, the Blue-gray Gnatcatcher, in
For example, although they may excavators, hawkers, aerial chasers, which he introduced the key
compete for the same food, on other and scavengers. concept of the guild, cemented
occasions they might cooperate to his reputation. Root was
deal with predators. Broad associations invited to join the staff of
Cornell University, where he
In the rush to identify guilds, there
taught biology and ecology.
was some confusion over just what While there, he researched
the term meant. By the 1980s, the the relationship between
American ecologists Charles arthropods (a large group of
Hawkins and James MacMahon invertebrates including insects
felt the need to redefine the term. and arachnids) and goldenrod
…does it matter that a They argued that the words “in a flowers. Root received many
particular insect species similar way” should be dropped awards during his career,
is captured by a silken from Root’s original definition. It including the Ecological
spider web as opposed does not matter, they maintained, Society of America’s Eminent
to a bird’s beak? whether an organism removes a Ecologist award in 2003
Charles Hawkins and tree leaf to build a nest or for food. and its Odum award in 2004.
James MacMahon It is the resource of the tree leaf
Key works
that matters rather than the way
it is utilized. Either way, the leaf- 1967 “The niche exploitation
users belong to a common guild pattern of the Blue-gray
because they are exploiting the Gnatcatcher”
same resource. ■
THE CITIZEN
NETWORK DEPENDS ON
VOLUNTEERS
CITIZEN SCIENCE
180 CITIZEN SCIENCE

North American migration pathways
IN CONTEXT
KEY FIGURES
Migratory birds
Fred Urquhart (1911–2002), in North America
Norah Urquhart (1918–2009) use paths that can
be divided into four
BEFORE north–south zones,
1883 The Bird Migration called flyways—Pacific,
and Distribution recording Central, Mississippi,
program starts in the US. and Atlantic. Citizen
scientists can play a
1966 The North American key role in recording
Breeding Birds Survey, the birds as they stop
KEY to feed or rest along
conducted by volunteers, the way, during their
begins in Maryland. Pacific flyway flights north in spring
and south in fall.
AFTER Central flyway
2007 The Global Biodiversity
Information Facility (GBIF) Mississippi flyway
launches a global online portal Atlantic flyway
for collecting data on plants
and animals from citizen
C
scientists and professionals. itizen science is research migration of birds, the Scottish
and observation carried enthusiasts using lighthouses
2010 The eBird online project,
out by nonprofessional around the coast as observation
created in the US in 2002 by individuals, teams, or networks posts. Then, in the early 1880s, the
the Cornell Laboratory of of volunteers, often in partnership idea of collective observation was
Ornithology for volunteers to with professional scientists. It is extended onto a national scale by
report real-time bird sightings, based on an appreciation that the American ornithologist Wells
becomes a global survey. scientific community should be Cooke, who began a project to show
responsive to the environmental arrival dates for migratory North
concerns of society as a whole, and American birds and provide
an understanding that citizens can evidence for migration pathways.
produce reliable scientific evidence Cooke’s project ran until World War
that leads to greater scientific II, gathering 6 million data cards
knowledge. The involvement of on more than 800 bird species and
ordinary people allows research utilizing 3,000 volunteers at its
bodies to accomplish projects that peak. In 2009, the North American
would be far too expensive or time- Bird Phenology Program began
consuming to run otherwise. to digitize the data from the cards,
Butterflies—millions which has provided valuable
upon millions … carpeted Early enthusiasts evidence of changed bird migration
the ground in their While the term “citizen science” dates and routes resulting from
flaming myriads on this is relatively new, dating from the global climate change.
Mexican mountainside. 1980s, the concept and practice The world’s longest-running
Fred Urqhuart of using the public to observe the citizen science survey is the
natural world and record data has Christmas Bird Count (CBC), held
a long pedigree. In the 1870s, each year in the US. Christmas
small groups of ornithologists “side hunts” of birds were a popular
in Germany and Scotland began pastime in many rural districts
to collect reports on the fall of the US in the 19th century,

See also: A system for identifying all nature’s organisms 86–87
■ Big ecology 153 ■ The distribution of species over space and time 162–163
regardless of whether the birds a tagging scheme in an attempt

were suitable for eating. In 1900, to find where the insect ended its
Frank Chapman, an officer of the journey after setting out from
Audubon Society—named after southern Canada and the northern
American ornithologist and painter states of the US in fall. They
John James Audubon—proposed enlisted the help of a small group
counting birds, rather than of “citizen scientists” to help tag
shooting them. He encouraged 27 the wings of the butterflies and
birdwatchers to participate in the report sightings. From a dozen or
first event, and the counts then so helpers, their Insect Migration
grew every year. In 2016–17, 73,153 Association, as it became known, Fred and
observers submitted counts from grew to hundreds of volunteers
Norah Urquhart
2,536 different locations in North who persisted for years, tagging
and Latin America, the Pacific, and hundreds of thousands of monarchs Born in 1911, Fred Urquhart
the Caribbean. The data on the with the message “Send to Zoology, grew up near a railroad line on
distribution and number of birds University of Toronto.” the edge of Toronto, Canada,
has provided a huge data set for Despite the Urquharts’ best and became intrigued by the
ecologists, allowing comparison efforts, the trail went cold in Texas. monarch butterflies that laid
over time and between habitats. Finally, on January 2, 1975, two their eggs close to the track.
amateur naturalists, Ken Brugger After graduating in 1937 from
In search of the monarch and Catalina Aguado, discovered the University of Toronto with
Perhaps the most celebrated act of the butterflies’ wintering site in bachelor’s and master’s
citizen science was one that set out montane forest north of Mexico ❯❯ degrees in biology, Urquhart
to solve the mystery of where the began to research the
butterfly. Having taught
migrating monarch butterfly went
Observations of birds made and meteorology to pilots during
in winter. In 1952, a Canadian recorded by “citizen scientists” in parks World War II, he returned to
couple, zoologists Fred and Norah and gardens can provide ecologists the university to lecture
Urquhart, who had long been with vital data on many species, such zoology and married Norah
fascinated by the butterfly, set up as the European Goldfinch. Roden Patterson, another
Toronto graduate, who joined
his quest to find the monarch’s
winter home. Fred Urquhart
also worked as Curator of
Insects and Director of
Zoology and Paleontology at
the Royal Ontario Museum.
In 1998, Fred and Norah
Urquhart were awarded their
nation’s highest civilian
award, the Order of Canada.
Key works
1960 The Monarch Butterfly

1987 The Monarch Butterfly:
International Traveler
182 CITIZEN SCIENCE
Science should be
dominated by amateurship
instead of money-biased
technical bureaucrats.
Erwin Chargaff
Austro-Hungarian biochemist
their own homes, but simply

to record what they saw in their
gardens, backyards, or streets.
By 2018, more than 500,000 people
were participating, recording
7 million birds. The vast amount
of data gathered can now be
compared for every year back
to 1979. Without public help, this
would simply not be possible.
In 1989, the term “citizen
science” first appeared in print,
in the journal American Birds. It
was used to describe a volunteer
project sponsored by the Audubon
Monarch butterflies form a cluster fall. Thousands of people Society that sampled rain for
to stay warm during migration. Tagging in Mexico, the US, and Canada acidity. The aim of the project
by volunteers revealed the monarch’s are helping build an ever clearer was to raise awareness of the
migratory routes, and continues with
the annual “Monarch Watch.”
picture of what routes the monarch acidification of rivers and lakes that
follows and how it deals with was killing fish and invertebrates,
changing weather patterns. and, indirectly, the birds that
City. No tagged monarchs were preyed on them. It was also
found, however, and it was not Citizens march on designed to put pressure on the
until the following January that the More volunteer-based projects US government, which soon after
Urquharts found one—tagged by were launched during the 1960s introduced the 1990 Clean Air Act.
two schoolboys in Minnesota the and 1970s, including the North Citizen science has also proved
previous August. Citizen science American Breeding Bird Survey, its worth for marine conservation.
had provided the hard evidence the British Nest Records Card In the Bahamas, a report in 2012
that the butterflies migrated from project, and a survey of sea turtle on declining numbers of the queen
North America to Mexico. Now egg laying in Japan. In 1979, the conch, a large sea snail, led to the
it is known where millions of Royal Society for the Protection of formation of “Conchservation,” a
monarchs spend the winter, the Birds (RSPB) launched the Big campaign that encourages locals
emphasis has changed to tracking Garden Birdwatch in the UK, which to tag conches. Another project,
their movements each spring and did not even require people to leave set up in the US in 2010, at the

University of Georgia, uses an Young volunteers at Siyeh Pass,
app, the Marine Debris Tracker, in the state of Montana, record their
to record sightings of debris in sightings of mountain goats for the
high country citizen science project
the ocean. Understanding patterns in the Glacier National Park.
of trash buildup in the world’s seas
helps scientists to track how
it is transported by currents and introduce bias into recordings,
where to concentrate removal efforts such as by the omission of a
for maximum effect. species that cannot be identified.
The advent of new technology Most simple citizen science
has led to a proliferation of citizen tasks, though, require no training,
science projects. Online recording and some other, more complex,
systems allow people to log procedures can be tackled after
sightings of anything from stag basic tuition. People are often
beetles to wildflowers or migrating attracted to citizen science precisely
birds. In the UK, for example, the because they gain new skills in the
Greenspace Information for Greater process. Increasing pressure on
London (GiGL) website, created by Earth’s natural environments and
the National Biodiversity Network, resources creates an ever greater Painting the
allows people to submit records need for data that records presence,
complete picture
online or by phone, adding to a absence, and change in species,
database used by scientists working their habitats, and the wider Citizen scientists are now
to conserve species and habitats. ecosystems. Projects such as the biggest global providers
Zooniverse, the world’s largest of data on the occurrence of
Limitations and potential citizen science platform, help living organisms. Data is
Some ecology research projects fill this need, accumulating data easier than ever to submit
are beyond the reach of untrained from around 1.7 million volunteers and artificial intelligence (AI)
amateurs because they require worldwide. Such projects will be an algorithms can process data
too high a degree of skill, or invaluable resource for conservation in minutes where once it
technology that is too complex organizations, research institutions, would have taken weeks. For
or expensive. People unfamiliar nongovernment agencies, and example, if a person records
with scientific methods may also governments for years to come. ■ sightings of birds coming to
a garden feeder and sends a
report from a phone to Cornell
University’s eBird website, the
information is compared with
Scientific study often relies on the collection
previous data on factors such
of large quantities of data.
as population numbers and
migration routes. More than
390,000 people have submitted
millions of bird sightings to
eBird from nearly 5 million
Volunteer networks are
The more data, the more able to collect vast
locations around the world.
representative the quantities of data, often
This data is fed into the Global
results are of reality. from widespread areas. Biodiversity Information
Facility (GBIF, coordinated in
Denmark), which collects
information on plants, animals,
fungi, and bacteria. GBIF now
The citizen’s network depends contains more than 1 billion
on volunteers. observations, and the number
is growing daily.
184
POPULATION DYNAMICS
BECOME CHAOTIC
WHEN THE RATE OF
REPRODUCTION SOARS
CHAOTIC POPULATION CHANGE
C
haos theory—the idea that
IN CONTEXT predictions are limited by
time and the nonlinear
KEY FIGURE
nature of behavior—took hold in the
Robert May (1936–)
1960s. American meteorologist
BEFORE Edward Lorenz observed the effect Chaos: when the present
1798 Thomas Malthus argues in weather patterns, and described determines the future, but
that human populations will it in 1961. Since then, the theory the approximate present
increase at an ever-faster rate, has been applied to many sciences, does not approximately
inevitably causing suffering. including population dynamics. determine the future.
Edward Lorenz
1845 Belgian demographist Chaotic populations
Pierre-François Verhulst argues In the 1970s, Australian scientist
that checks to population Robert May became interested in
growth will increase in line animal population dynamics, and
with population growth itself. worked on a model to forecast
growth or decline over time. This
AFTER led him to the logistic equation. patterns at the lowest rates of
1987 Per Bak, Chao Tang, and Devised by Belgian mathematician growth, May found that the logistic
Kurt Wiesenfeld, a research Pierre-François Verhulst, this equation produced erratic results
team in New York, describe equation produces an S-shaped when the growth rate was equal to
“self-organized criticality”— curve on a graph—showing or above 3.9. Instead of producing
elements within a system population growing slowly at first, repeating patterns, the map
interacting spontaneously then rapidly, before tapering off plotted trajectories that appeared
to produce change. into a state of equilibrium. completely random. May’s work
May experimented with showed how a simple, constant
2014 Japanese ecologist Verhulst’s formula to create the equation could produce chaotic
George Sugihari uses a chaos “logistic map,” which showed behavior. His logistic map is now
theory approach called empirical the population trends on a graph. used by demographers to track and
dynamic modeling to produce Although it created predictable predict population growth. ■
a more accurate estimate of
salmon numbers in Canada’s See also: Predator–prey equations 44–49 ■ Non-consumptive effects of predators
Fraser River. on their prey 76–77 ■ The Verhulst equation 164–165 ■ Metapopulations 186–187
TO VISUALIZE
THE BIG PICTURE,
TAKE A
MACROECOLOGY
DISTANT VIEW
S
cientists seeking faster mammal species, Brown was able
IN CONTEXT ways to analyze and counter to work out the extinction risk on
the many threats to plant each ridge as temperatures rose,
KEY FIGURE
and animal populations increasingly and suggest conservation priorities.
James H. Brown (1942–)
turn to macroecology. The term,
BEFORE coined by American ecologists Enhancing fieldwork
1920 Swedish ecologist James Brown and Brian Maurer Macroecology often supplements
Olof Arrhenius produces in 1989, describes studies that fieldwork and can lead to surprising
a mathematical formula for examine relationships between discoveries. In Madagascar,
the relationship between organisms and their environment satellite data was used to develop
area and species diversity. across large areas to explain models for chameleon species and
patterns of abundance, diversity, predict them in areas beyond
1964 British entomologist distribution, and change. their known ranges. As a result,
C.B. Williams documents Brown had tried and tested this scientists investigating these areas
patterns of species abundance, methodology in the 1970s while found several new sister species. ■
distribution, and diversity in studying the potential effects of
his book Patterns in the global warming on species in cool,
Balance of Nature. moist forest and meadow habitats
on 19 isolated ridges of the Great
AFTER Basin, in California and Utah. He
2002 British ecologists Tim realized it would take years of fresh
Blackburn and Kevin Gaston fieldwork to collect enough data.
argue—contrary to some— Instead, he used existing findings
that macroecology should be to draw new conclusions. First, he
treated as a discipline distinct predicted how much shrinkage
from biogeography. would occur in the area of ridge-top
By comparing community studies
habitat with an assumed increase made in deserts around the world,
2018 A team of scientists uses in temperature. Using known data macroecologists can determine the
practical macroecological on the minimum area required to greatest threats to a desert species
methods to show that bird support a population of each small such as this banner-tailed kangaroo rat.
species living on islands have
relatively larger brains than See also: Field experiments 54–55 ■ Animal ecology 106–113 ■ Island
their mainland relatives. biogeography 144–149 ■ Big ecology 153 ■ Endangered habitats 236–239
186
A POPULATION
OF POPULATIONS
METAPOPULATIONS
IN CONTEXT A species becomes

A species colonizes
extinct in one
KEY FIGURE an empty habitat patch.
habitat patch.
Ilkka Hanski (1953–2016)
BEFORE
1931 In the US, geneticist
Sewall Wright explores the
influence of genetic factors Extinction and colonization are dynamic processes.
on species populations.
1933 In Australia, ecologist
Alexander Nicholson and
physicist Victor Bailey develop A local extinction does not signal
their model of population the extinction of the species.
dynamics to describe the
host–parasite relationship.
A
1954 In The Distribution and metapopulation is a other places. The species is like a
Abundance of Animals, combination of separate, family whose members have moved
Australian ecologists Herbert local populations of the to different cities yet are still
Andrewartha and Charles same species. The term was coined related. The combined effect of
Birch challenge the idea that by American ecologist Richard many populations may boost the
species populations are Levins in 1969 to describe how long-term survival of the species.
controlled by density alone. insect pest populations rise and fall
on farm fields. Since then, its use Apart but together
AFTER has expanded to cover any species A crucial aspect of metapopulation
2007 American ecologist broken up into local populations in theory is the level of interaction
James Petranka links fragmented habitats, both on land between the separate local
metapopulation theory to and in the oceans. populations. If the level is high,
the metamorphosis stages A particular species of bird, for it is not considered to be a
of amphibians. instance, may be found in separate metapopulation—all the local
populations in a lowland forest, in groups are part of one big
mountain woodlands, and various population. In a metapopulation

See also: Animal ecology 106–113 ■ Clutch control 114–115
■ Island biogeography 144–149 ■ Metacommunities 190–193
contact between the various local leaving vacant patches for another
groups is limited, and they remain population to recolonize. Hansk
partly cut off in their own local argued that there is persistent
habitat or “patch.” Yet there has to balance between “deaths” (local
be at least some interaction. It may extinctions) and “births” (the
be just a single brave or outcast establishment of new populations
member of one group that enters at unoccupied sites). He likened
another patch and mates with the this balance to the spread of
local population there. Isolation for disease, with the susceptible and
too long pushes local populations the infected representing in turn
apart to the point where they can empty and occupied “patches” Ilkka Hanski
no longer mate with one another, for disease-carrying parasites.
and in time they become separate Ecologists see the concept of Widely seen as the father of
species or subspecies. metapopulations as increasingly metapopulation theory, Ilkka
In the 1990s, Finnish ecologist important in understanding how Hanski was born in Lempäälä,
Ilkka Hanski showed that at the species will survive, particularly Finland, in 1953. As a child, he
core of metapopulation theory is the in the face of human influence on collected butterflies, and after
notion that local populations are habitats. The theory helps them finding a rare species, he
unstable. The metapopulation as analyze the way populations rise devoted his life to ecology,
a whole may well be stable, but the and fall, using mathematical models studying at the universities
local populations are likely to rise to play out interactions, and enables of Helsinki and Oxford.
and fall in their individual patches them to predict how much habitat Ecologists at the time
paid little attention to the
in response to inside and outside fragmentation a species can endure
distribution of local species
influences. Some patch members before it is driven to extinction. ■
populations, but Hanski
may emigrate and join a much realized this was crucial,
reduced population in danger and spent much of his career
The Glanville fritillary butterfly
of extinction, giving it renewed metapopulation, in its fragmented testing his metapopulation
strength—a metapopulation feature habitats on Finland’s Åland Islands, theory by mapping out and
known as the “rescue effect.” Other provided the ideal subject for Ilkka recording more than 4,000
groups may completely vanish, Hanski’s studies into species patches. habitat patches for the
Glanville fritillary butterfly
on the Åland Islands. This
work earned Hanski global
fame, and enabled him to
establish the Metapopulation
Research Centre in Helsinki,
which became one the world’s
leading focuses of ecological
research. Hanski died of
cancer in May 2016.
Key works
1991 Metapopulation
Dynamics
1999 Metapopulation Ecology
2016 Messages from Islands
188
ORGANISMS CHANGE AND

CONSTRUCT THE WORLD
IN WHICH THEY LIVE
NICHE CONSTRUCTION
A
ll organisms alter the He argued that they actively
IN CONTEXT environment to cater to construct and modify their
their own needs. Animals environment, and affect their own
KEY FIGURE
dig burrows, build nests, create evolution in the process: the lynx
F. John Odling-Smee
shade from the sun, and create and the hare, for example, shape
(1935–)
shelter from the wind to provide a each other's evolution and shared
BEFORE more secure environment, while environment by striving to outrun
1969 British biologist Conrad plants alter soil chemistry and cycle each other. Odling-Smee similarly
Waddington writes about nutrients. When organisms modify argued that niche construction and
ways in which animals change their own and each other’s place “ecological inheritance”—when
their environments, calling in the environment, this is “niche inherited resources and conditions
this “the exploitive system.” construction”—a term coined such as altered soil chemistry are
by British evolutionary biologist passed on to descendants—should
1983 Richard Lewontin, an F. John Odling-Smee in 1988. be seen as evolutionary processes.
American biologist, argues American evolutionary biologist
that organisms are active Richard Lewontin had previously Levels of construction
constructors of their own suggested that animals are not Some common examples of niche
environments, in Gene, passive victims of natural selection. construction are obvious, while
Organism, and Environment. others operate at a microscopic
scale. Beavers build impressive
AFTER dams across rivers, creating lakes
2014 Canadian ecologist Blake and altering river courses. This
Matthews outlines criteria for alters the composition of the water
deciding whether an organism and materials carried downstream,
is a niche constructor. Hares do not sit around creates new habitats for other
constructing lynxes! But organisms to take advantage of,
in the most important and also changes the composition
sense, they do. of the river’s plant and animal
Richard Lewontin communities. British biologist
Kevin Laland has suggested that,
while a beaver’s dam is clearly of
great evolutionary and ecological
importance, the impact of its dung
may also be significant.

See also: Ecological niches 50–51 ■ The ecosystem 134–137 Organisms and
their environment 166 ■ The ecological guild 176–177
■
Ecosystem engineers
Niche constructors have been
described as “ecosystem
engineers,” a term coined in
1994 by scientists Clive Jones,
John Lawton, and Moshe
Shachak. They outlined two
kinds of ecosystem engineers.
The first, allogenic ecosystem
engineers, change physical
materials. Take, for example,
beavers building dams,
woodpeckers excavating
nest holes, and people mining
for gravel; these activities
modify the availability of
resources for other species.
When woodpeckers abandon
their holes, smaller birds and
other animals move in. If water
floods a gravel pit, ducks and
dragonflies can colonize it.
Other ecosystem engineers
are autogenic, which means
that simply by growing, they
provide new habitats for other
plants and animals. A mature
Earthworms leave castings that British biologists Nancy Harrison oak tree, for example, is a
make them valuable natural fertilizers. and Michael Whitehouse have also suitable environment for a
They not only transform the soil for suggested that when birds form broader range of insects, birds,
themselves but also help plants to grow. and small mammals than an
mixed-species flocks—as many do
outside of the breeding season— oak sapling. Likewise, a coral
Earthworms are highly effective they are altering their relationship reef provides homes for more
fish and crustaceans as it
niche constructors, constantly with competitors to find more food
grows larger.
transforming the soil in which they resources and gain more protection
live. They break down vegetable from predators. The complex social
and mineral matter into particles environment they create modifies
small enough for plants to ingest. their own ecology and behavior.
The worm casts they secrete are In his explanation of niche
five times richer in usable nitrogen, construction, Odling-Smee pointed
have seven times the concentration to ancient cyanobacteria, which
of phosphates, and are about 11 produced oxygen as a by-product of
times richer in potassium than the photosynthesis more than 2 billion
surrounding soil. years ago. This was a key factor
Similarly, microscopic diatoms in the Great Oxygenation Event,
living in seafloor sediments secrete which changed the composition
chemicals that bind and stabilize of Earth’s atmosphere and oceans,
the sand. In Canada’s Bay of Fundy, massively modifying our planet's
for example, the changes diatoms environment. The oxygen boost A European Starling in Arizona,
US, takes advantage of a hole
make to the physical state of the helped create the conditions for the abandoned by a Gila Woodpecker
seabed allow other organisms, such evolution of much more complex life to make its own nest.
as mud shrimp, to colonize it. forms—including humans. ■
190
IN CONTEXT
LOCAL COMMUNITIES KEY FIGURE

Mathew Leibold (1956–)
THAT EXCHANGE
BEFORE
1917 Arthur Tansley observes
that two species of Galium
COLONISTS
plants grow differently in
different soil patches.
METACOMMUNITIES 1934 Georgy Gause develops

the competitive exclusion
principle stating that two
species competing for the
same key resource cannot
coexist for long.
2001 Stephen Hubbell’s
“neutral theory” argues that
biodiversity arises at random.
AFTER
2006 Mathew Leibold and
fellow American ecologist
Marcel Holyoak refine and
develop the theory of
metacommunities.
O
ne of the limitations
of traditional community
ecology was that it tended
to look at communities purely
locally and take little account of
what happens at different scales
or across different places. Therefore,
over the last few decades, ecologists
have been developing theories of
“meta” communities; the concept
was summed up in 2004 in a key
paper led by American ecologist
Mathew Leibold.
The idea of metacommunities
is linked to that of metapopulations.
While studies of metapopulations
examine the different patches
where populations of the same
species coexist, in metacommunity

See also: Competitive exclusion principle 52–53 ■ The ecosystem 134–137 ■ The neutral theory of biodiversity 152
■ Metapopulations 186–187
Mountain goats in Colorado live

in a metacommunity of species in
a mountain range—the Rocky
Mountains—but within a population
of goats on one single peak.
theory the different patches consist

of entire communities that include
a number of interacting species.
What is a metacommunity?
Metacommunities are essentially
groups or sets of communities.
The communities making up a
metacommunity are separated in
space, but they are not completely
isolated and independent. They feed, shelter, or breed. Differing of seemingly contradictory
interact as various species move types of habitat will influence this observations. One ecologist’s study,
between them. For example, a balance between interlinked and for instance, might look at the way
metacommunity might consist of a independent development. The species live and interact together
set of separate forest communities, theory of metacommunities provides in a small local community. This
spread across a region. The various a framework for studying how and narrowly focused study finds that
species within each patch of forest why variations develop and their competition between species for
habitat interact as an independent impact on biodiversity and resources is a crucial factor in the
community. However, certain population fluctuations. workings of the community.
species, including deer or rabbits, Another study might look at the
may migrate or disperse to another Local versus regional picture across a larger community.
community in the metacommunity, A major advantage of looking at This macro-study discovers that
moving to a different patch of forest communities in this spatial way competition plays virtually no
in search of better opportunities to is that it may help resolve a number part. So which result is correct? ❯❯
Wildlife crossings Many different species cross from one habitat to another. The
naturally between separate idea of providing wildlife with
habitat patches. This movement ways through is not new. For
can be seasonal, as in annual example, fishways for fish to
migrations, or prompted by bypass dams go back centuries.
natural disasters, such as fire or Wildlife crossings—from bridges
flood, or may take place over long for bears in Canada to tunnels
timespans. It creates connections for California’s desert tortoises—
that are often essential for the are becoming an increasingly
health and survival of species and common feature of construction
communities, providing renewal work. Thousands of crossings,
or new resources at pivotal among them bridges, viaducts,
moments. Increasingly, however, and underpasses—often planted
manmade barriers, such as with vegetation—have been
clearances for agriculture, road, built to conserve habitats and
railroads, and urban sprawl, are to avoid fatal collisions between
breaking up this natural interflow animals and vehicles.
192 METACOMMUNITIES
The answer may be that both are conflicting theories. It may make stochastic processes, such as the
right, and the difference simply it easier, for example, to resolve spread of a species by freak storms
depends on scale. The benefit of the century-old debate between the or a die-off due to an epidemic.
metacommunity theory is that it “deterministic,” niche-based theory It also acknowledges that regional
allows ecologists to reconcile these of community ecology, in which changes can be caused by the
differences. It enables them to look species diversity is determined combined effect of local ones.
for explanations on both a local and by each species’ ecological
regional scale. niche, and “stochastic” (random) Finding metacommunities
A metacommunity might be a theory, which emphasizes the One of the problems with Leibold’s
set of half a dozen deciduous trees importance of chance colonization concept is that in practice it is
within a park, with each tree an and ecological drift (random not so straightforward to identify
individual community. However, fluctuations in population sizes). the separate components of a
it could equally be all the deciduous Metacommunity theory provides metacommunity. For the fish and
forests in temperate zones all around an umbrella framework for seeing other water creatures in different
the world. What metacommunity how deterministic and stochastic lakes within a lake district, for
theory does is allow ecologists to processes can interact to form instance, each lake may clearly be
work at any scale, at least in theory. natural communities. It allows a distinct community. However, for
ecologists to state that patterns those birds able to fly between the
Umbrella framework of biodiversity are determined both lakes in minutes, the different lakes
According to Mathew Leibold, the by local biological features, such are all part of the same single
study of metacommunities brings as the balance of sun and shade in community. This may explain why
together many seemingly disparate rock pools or variations in water much of the continuing work and
branches of ecology and apparently quality in streams, and by regional research on metacommunities
Metacommunity
perch dragonfly pond weed

dragonfly
lake 2
lake 1
frog
heron cormorant
rushes heron frog
lake 3
perch mosquito
rushes algae carp cormorant
lake 4 lake 5
mosquito
duck algae
mayfly
In this example of a metacommunity, arrows show

how species move between lakes to feed or breed. Seeds
and the spores of algae are dispersed by the wind.

Rockpools in a wave-cut platform
form a metacommunity on Eysturoy
in the Faroe Islands. The rockpools are
separate between tides, but become
joined as one when the tide comes in.
for a period of time following a

storm, patches of fungal fruiting
bodies that live just a few days
or weeks, and even pitcher plants
that, after dew or rain, provide a
short-lived aquatic home for both
bacteria and insects.
Blurred communities
Leibold’s 2004 paper acknowledged
that the metacommunities with
blurred boundaries are perhaps the
hardest to define. Coral reefs, for
example, may look neatly separate,
but many of the species that live
among them swim freely and
respond to a host of changing
outside influences, such as
shifts in ocean currents.
has been theoretical and abstract study and there is a vast literature Since most of the world’s life
rather than rooted in fieldwork. on island biogeography, reaching exists within such vaguely defined
Some metacommunities are easy back to Charles Darwin’s famous patches, theorists have attempted
to identify, such as islands in an study of variations between finches further clarification. Leibold and
island group, or rockpools that are in the Galapagos Islands in the his colleagues have suggested
separate between tides but joined Pacific Ocean. Neatly separate two different ways of identifying
when the tide comes in. In their patches make good subjects for metacommunities for study:
2004 paper, Leibold and his study, which is why they have been distinct communities embedded
colleagues acknowledged that local popular with community ecologists. within a “matrix” habitat, such
communities, or patches, do not But, of course, birds and many as clearings in a forest rich in
always have clear boundaries that other organisms blown across by resources; and arbitrary sampling
make them recognizably separate, the wind or washed in by the sea patches in a continuous habitat,
and that different species may ensure that even island communities such as a random circle of trees
respond to things happening at a are never completely isolated. This within a forest.
different scale. They identified three is why some metacommunity The work is still at an early
kinds of metacommunity: markedly studies focus on the space between stage. The world is entering a
separate patches; short-lived but the communities even where the biodiversity crisis, and countless
distinct patches that appear in a patches are distinct, as they are species and communities appear
habitat from time to time at varying with ponds and lakes, and analyse to be under threat from the effects
size; and permanent patches with how species move between them. of human activity. Metacommunity
vague or “blurred” boundaries. Short-lived but distinct patches theory may, in time, help to provide
may be much harder to identify, a better understanding of how
Distinct patches simply because of their ephemeral natural communities will respond,
The most obvious markedly separate nature. Nonetheless, ecologists and how local changes to habitats
patches are islands in the ocean. have made metacommunity studies may ripple through a region, either
These are a convenient subject to of holes in trees that fill with water adversely or positively. ■
THE LIVI
EARTH
NG
196 INTRODUCTION
Louis Agassiz shows that Vladimir Vernadsky’s

an ice sheet once covered Svante Arrhenius is the book The Biosphere explains
Switzerland, and suggests first to argue that carbon how atmospheric gases are
that an ice age occurred dioxide emissions can lead created by biological
in recent geological history. to global warming. processes.
1840 1896 1926
1869 1912 1935
The father of biogeography Alfred Wegener presents his Arthur Tansley coins the term
Alfred Russel Wallace theory that Earth was once “ecosystem” to describe
reports a clear evolutionary a single landmass from an interdependent
division in fauna species which continents community of biological and
on neighboring islands. drifted apart. nonbiological components.
F
or centuries, scientists must indicate a truly ancient origin continents were once joined and
in the Western world tried for the planet—an idea which Lyell had broken away. It was not until
to reconcile the findings developed further in the 1830s. the 1960s that a viable mechanism
of geologists and fossil hunters Soon after, Swiss-American was found for such movement.
with literal interpretations of geologist Louis Agassiz proposed Geophysicists discovered patterns
biblical stories about Creation that the topography of some regions of magnetic anomalies running in
and the Great Flood. In 1654, for had been shaped by glaciations. parallel stripes on either side of
example, Archbishop Ussher dated Hutton and Lyell also noted that ocean ridges and identified the
Earth’s creation to October 22, fossils of animals and plants process of seafloor spreading—hot
4004 BCE. A series of discoveries vanished from the geological magma bubbling up through cracks
challenged this narrative and led record. Lyell believed this to be in the oceanic crust and forming
to new ideas about the dynamic evidence of extinction, challenging new crust as it cools and moves
history of life on Earth. the prevailing belief that species away. This gradual process shifts
were immutable. and shapes continents.
Evidence in the rocks Fossils also offered clues to
Two Scottish geologists—James movements of Earth’s continents. The birth of biogeography
Hutton and Charles Lyell— German meteorologist Alfred In the Age of Exploration from the
advanced our understanding of Wegener noted that similar fossils 16th century on, scientists began to
Earth’s age. In Theory of the Earth could be found on both sides of the study the geographical distribution
(1795), Hutton argued that the South Atlantic, even though they of plants and animals. By the 1860s,
repeated cycles of sedimentation were thousands of miles apart. In Alfred Russel Wallace viewed these
and erosion necessary to create his 1912 theory of continental drift, patterns, clearly defined by physical
thousands of feet of rock strata Wegener cited this as evidence that barriers such as mountains and
THE LIVING EARTH 197
In their book Bioecology, Eugene and Howard Odum James Lovelock’s Gaia
Frederic Clements and describe the living planet hypothesis presents
Victor Shelford popularize as a global collection of Earth as a single,
the idea of the biome. interlocking systems. self-regulating system.
1939 1953 1974
1947 1970 1980
Lesley Holdridge’s The first Earth Day is Luis and Walter

biogeographic classification celebrated, and the idea Alvarez suggest that the
maps 38 life zones where that humans should view mass extinction of the
vegetation is determined Earth holistically dinosaurs was caused
by rain and heat. grows in popularity. by a meteor strike.
seas as a key supporting argument areas at or near the surface of the Almost two centuries earlier Hutton
for evolution. Wallace noted, for Earth where organic life can exist. had articulated a similar idea—that
example, the ocean straits that In 1926, the Russian geochemist biological and geological processes
produced a sharp division between Vladimir Vernadsky explained are interlinked and that Earth could
the flora and fauna of Australasia the biosphere’s close interaction be viewed as a superorganism. In
and Southeast Asia. with the planet’s rock (lithosphere), Hutton’s words, “The globe of this
With a better understanding of water (hydrosphere) and air earth is not just a machine but also
Earth’s biogeography, 20th-century (atmosphere). This in turn led an organized body as it has a
ecologists divided the planet into American biologist Eugene Odum regenerative power.”
biomes—broad communities of flora to advocate a holistic approach to
and fauna that interact in different ecology. Odum argued that it was Heading for extinction?
habitats, such as tropical rain forests not possible to understand a single Life has survived on Earth for
or tundra. Botanist Leslie Holdridge organism, or a group of organisms, billions of years, despite the ravages
refined the concept in 1947 with his without studying the ecosystem in of five mass extinctions. However,
life zone classification, in which he which they live. He described this environmentalists now question
mapped zones based on the two view as “the new ecology.” whether it will survive another.
crucial influences on vegetation: In 1974, British scientist James Indeed, some contend that a sixth
temperature and rainfall. Lovelock advanced the Gaia mass extinction has already started,
hypothesis that the interaction as a result of human activity. Yet,
A “whole Earth” approach of living and nonliving elements if Lovelock’s Gaia theory is correct,
The word “biosphere” was coined in the biosphere reveal Earth to be a it seems likely that the planet will
by Austrian geologist Edward complex, self-regulating system that endure—even if humans and many
Suess in 1875 to signify all the perpetuates the conditions for life. other current life forms do not. ■
198
THE GLACIER
WAS GOD’S
GREAT PLOW
ANCIENT ICE AGES
I
n the early 19th century, there Earth’s history has been a process
IN CONTEXT were contradictory explanations of slow change, punctuated by
for the development of Earth’s catastrophic events. The study
KEY FIGURE
landforms, plants, and animals. of glaciers, and the landforms they
Louis Agassiz (1807–73)
Supporters of catastrophism argued create, informed these ideas. After
BEFORE that a series of destructive shocks, observing parallel striations in rocks
1795 Scottish geologist James such as the Great Flood described in of the Swiss Alps, German–Swiss
Hutton argues that erratic the Bible, had re-formed the surface geologist Jean de Charpentier (or
boulders (rock fragments that of the planet many times, reshaping Johann von Charpentier) postulated
are different from the existing mountains, lakes, and rivers that glaciers in the Alps had once
underlying rock) in the and wiping out many plant and been more extensive and had
Alps were transported by animal species. In contrast, followers caused the scratches as they moved
moving glaciers. of uniformitarianism contended that and their sediment cut into the rock.
Earth’s features were the result of Geologist Jens Esmark drew similar
1818 In Sweden, naturalist continuous and uniform natural conclusions in Norway.
Göran Wahlenburg publishes processes of erosion, sedimentation
his theory that ice once (the depositing of particles carried Glacier movements
covered Scandinavia. by fluid flows), and volcanism. Swiss zoologist Louis Agassiz
Detailed geological studies developed Charpentier’s and
1824 Danish–Norwegian demonstrated that neither camp Esmark’s ideas further. In 1837,
mineralogist Jens Esmark was right. They established that he proposed that vast sheets of
theorizes that glaciers were ice had once covered much of the
once larger and thicker and northern hemisphere, from the
had covered much of Norway North Pole to the Mediterranean and
and the adjacent seafloor. Caspian coastlines. Agassiz also
undertook some detailed studies
AFTER of glacier movement in Switzerland
1938 Serbian mathematician and published his Études sur les
Milutin Milanković publishes
a theory to explain the
recurrence of ice ages based Animals enter Noah’s ark in a
depiction of the Great Flood described
on changes in Earth’s orbit in the Bible. Catastrophists believed that
around the Sun. the Great Flood was one of the formative
shocks that shaped the geology of Earth.

See also: Evolution by natural selection 24–31 ■ Global warming 202–203
■ The Keeling Curve 240–241 ■ Ozone depletion 260–261 ■ Spring creep 274–279
Receding glaciers
and bird migration
When the last glacial period
began to end, around 26,500
years ago, Earth was much
colder than it is today. Much
of North America and northern
Eurasia was covered with ice
sheets. The environment was
so harsh that most birds
tended to live in subtropical
and tropical regions where
there was more food.
As temperatures began
rising, the ice sheets started
to shrink, uncovering a new
landscape. Ice-free ground
and short, wet summers were
ideal for insects, and birds
began to move in, too, to
take advantage of this food
supply. When days got shorter
in fall, some birds stayed on
for the winter, but others
returned south.
glaciers in 1840. The same year, he Glaciers converge on Piz Argient, The distances flown by
visited geologist William Buckland a mountain in the Swiss Alps. Like birds returning to their homes
others in the Alps, these glaciers were
in Scotland to investigate glacial grew longer as the ice sheets
once much more extensive than they
features there, prompting Scottish are now, and they continue to shrink. retreated farther, eventually
glaciologist James Forbes to begin developing into long-distance
similar research in the French Alps. spring and fall migrations
Some quarters, such as the Aerial surveys in the 1920s and between the tropics and
Catholic Church, still argued that 1930s confirmed the extent of their northern latitudes. Common
glacial striations had been caused vast ice sheets—now defined birds that undertake the
by a great flood or that large silt and as areas of glacier ice exceeding journey include swallows,
warblers, and cuckoos.
rock deposits had been transported 19,300 sq miles (50,000 sq km); ice
by icebergs swept along by the flood. caps, such as Iceland’s Vatnajökull,
From the 1860s, however, there was are smaller.
wide support for Agassiz’s glaciation Further evidence revealed that
theory and the idea that glaciers in there had not been one single ice
the Swiss Alps and Norway had age, but at least five major ice ages
once extended much further. It was in Earth’s long history. The most
also accepted that a sheet of ice recent, the Quaternary Ice Age,
had once spread across Europe, and began 2.58 million years ago and is
south from the Arctic through much ongoing. In the last 750,000 years,
of North America, with catastrophic there have been eight ice advances
implications for plants and animals. (glacial periods) and retreats
By the late 1800s and early (interglacial periods). During the last A male Baltimore oriole perches
1900s, as more expeditions to both glacial period, which ended 10,000– on a tree fern in Costa Rica. The
species flies north to breed in
Greenland and Antarctica were 15,000 years ago, ice sheets were up March and returns to the tropics
undertaken, it became known that to 2½ miles (4 km) thick, and the sea in August or September.
both areas were still covered in ice. level was 390 ft (120 m) lower. ■
200
THERE IS NOTHING
ON THE MAP TO MARK
THE BOUNDARY
BIOGEOGRAPHY
LINE
T
he places where animals In the 18th century, as explorers
IN CONTEXT and plants live often vary recorded the plants and animals
in a regular manner along they saw, a picture of geographic
KEY FIGURE
geographic gradients of latitude, change had begun to emerge. On
Alfred Russel Wallace
elevation, and habitat type. The the great 1831–36 expedition of
(1823–1913)
study of this variation is known HMS Beagle, Charles Darwin saw
BEFORE as biogeography. One branch species of birds on the Falkland
1831–36 Darwin’s studies (phytogeography) examines the Islands that did not live on
on the voyage of HMS Beagle distribution of plants, whereas mainland South America, giant
confirm that many animals the other (zoogeography) analyzes tortoises that were unique to the
living in one area are not found the distribution of animals. British Galapagos Islands, and marsupials
in similar habitats elsewhere. naturalist and biologist Alfred such as Australia’s kangaroos. New
Russel Wallace is widely regarded pieces of the biogeographic jigsaw
AFTER as the “father of biogeography.” were falling into place.
1874 British zoologist Philip
Sclater categorizes birds by
zoogeographic regions. Zoogeographic regions of the world
1876 Alfred Russel Wallace
publishes The Geographical
Distribution of Animals—the
first extensive publication N EA RC TIC PALEARC TIC
on biogeography.
1975 Hungarian biogeographer IND OMALAYA
Miklos Udvardy proposes
WALLACE’S LINE
dividing biogeographic realms AFROTROPIC S
into biogeographic provinces. NEOTROPIC S
AU S TRALAS IA
2015 Mexican evolutionary
biologist J.J. Morrone proposes
an International Code of Area
Nomenclature for biogeography. Wallace’s six zoogeographic regions began with the
line he proposed in 1859 to mark the division of fauna
between Southeast Asia and Australasia.

See also: Evolution by natural selection 24–31 ■ Island biogeography 144–149
■ The distribution of species over space and time 162–163 ■ Biomes 206–209
Alfred Russel Wallace

Explorer, naturalist, biologist,
geographer, and social
reformer Alfred Russel
Wallace left school at 14,
From 1848, Wallace conducted All of Siberia is in the Palearctic and trained as a surveyor
years of fieldwork in South America region, and the Siberian in London before becoming
and Southeast Asia. He researched white birch trees depicted here are a teacher. He became
the feeding and breeding behavior part of a subdivision called the East fascinated with insects after
Siberian taiga. meeting British entomologist
and migratory habits of thousands
of species, paying specific attention Henry Bates. The pair
to animal distribution compared Bali and Lombok); this separates ventured to the Amazon
with the presence or absence of Asian fauna from the Australasian. Basin in 1848 on a four-year
collecting expedition. Trips to
geographical barriers, such as seas He found that larger mammals and
the Orinoco River and the
between islands. He concluded that most birds did not cross the line. Malay Archipelago followed.
the number of organisms living in a For example, tigers and rhinos live Wallace arrived at the same
community depends on the food only on the Asian side; babirusas, conclusion as Charles Darwin
available in that specific habitat. marsupials, and sulfur-breasted on the origin of species by
cockatoos only on the other side. natural selection, and they
Wallace’s Line He also highlighted the sharp presented their papers jointly
During his 1854–62 expedition differences between animals in in 1858. A world authority on
to the Malay Archipelago, Wallace North and South America. fauna distribution, Wallace
collected an astonishing 126,000 In 1876, Wallace proposed six also raised awareness about
specimens, many of them from separate zoogeographic regions: problems caused by human
species unknown to Western Nearctic (North America); Neotropics impact on the environment.
science, including 2 percent of the (South America); Palaearctic
world’s bird species. He regarded (Europe, Africa north of the Sahara Key works
biogeography as support for the Desert, and Central, North, and East
theory of evolution by means of Asia); Afrotropics (Africa south of 1869 The Malay Archipelago
1870 Contributions to the
natural selection. One of Wallace’s the Sahara Desert); Indomalaya
Theory of Natural Selection
important findings was the marked (South and Southeast Asia); and
1876 The Geographical
difference in bird species on either Australasia (Australia, New Guinea, Distribution of Animals
side of what was to become known and New Zealand). Today Wallace’s 1878 Tropical Nature, and
as the Wallace Line, which runs regions, with the addition of Other Essays
along the Makassar Strait (between Oceania (the islands of the Pacific 1880 Island Life
the islands of Borneo and Sulawesi) Ocean) and Antarctica, are known
and the Lombok Strait (between as biogeographic realms. ■
202
GLOBAL WARMING
ISN'T A PREDICTION.
IT IS HAPPENING.
GLOBAL WARMING
I
n 1896, Swedish chemist “greenhouse gases,” as they are
IN CONTEXT Svante Arrhenius became now known, and believed that
the first person to argue that increasing levels of CO2 would
KEY FIGURE
carbon dioxide (CO2) emissions raise Earth’s temperature. More
Svante Arrhenius
caused by human beings could specifically, he estimated that if
(1859–1927)
lead to global warming. Arrhenius levels of carbon dioxide increased
BEFORE thought that the average ground by 2.5 to 3 times, Arctic regions of
1824 French physicist Joseph temperature could be influenced the world would see temperature
Fourier suggests that Earth’s by carbon dioxide and other increases of 14–16°F (8–9°C).
atmosphere traps the Sun’s
heat like a greenhouse. The greenhouse effect
1859 Irish physicist John
Water vapor and other gases in Earth’s
Tyndall provides experimental atmosphere, such as carbon dioxide and
evidence to support earlier SUN methane, trap heat from the Sun and
hypotheses that atmospheric infrared radiation from Earth, raising
gases absorb radiant heat. the planet’s temperature.
AFTER
1976 American scientist
Some heat escapes
Charles Keeling proves that
Sol
into space
between 1959 and 1971
ar r
carbon dioxide levels in the

ad i
atmosphere increased by Some heat is trapped

atio
about 3.4 percent each year. in the atmosphere

RE
n
n
r ad i a re d
E
atio
2006 In Field Notes from PH

S
Infr
a Catastrophe, journalist
O
M
Elizabeth Kolbert tells the

AT
stories of people and places

H ’S
impacted by climate change.

EART
EARTH

See also: Environmental feedback loops 224–225 ■ Renewable energy 300–305
■ The Green Movement 308–309 ■ Halting climate change 316–321
The effects of
global warming
Arrhenius was building on the Since the end of the 19th

work of scientists Joseph Fourier century, carbon dioxide (CO2)
in the atmosphere has
and John Tyndall earlier in the
increased by about 25 percent,
19th century. Fourier had wondered and the average global
why Earth was not a freezing temperature by around 0.9°F
wasteland, when the Sun was too The atmosphere may act like (0.5°C). Scientific evidence
far away to heat it to its current the glass of a greenhouse … proves that these changes
temperature. He knew that heated [raising] the mean temperature have contributed to melting
surfaces—such as the surface of of Earth’s surface. glaciers and sea ice followed
Earth—emit thermal energy, and Nils Ekholm by rising sea levels—around
that the thermal energy radiating Swedish meteorologist (1848–1923) 8 in (20 cm) since 1880—as
back into space should result in well as damage to coral reefs.
colder temperatures on Earth. Other phenomena include
Something was regulating the longer wildfire seasons, more
temperature, and Fourier theorized extreme weather, and shifts in
that Earth’s atmosphere, made the ranges of animals and
up of various gases, acted like a plants, leading to disease,
glass box, containing the air average 59°F (15°C), although in extinction, and food shortages.
The extent to which global
and keeping it warm. Fourier’s recent decades human activities
temperatures will increase
hypothesis, although oversimplistic, that release greenhouse gases
depends on whether (and
led to the “greenhouse effect” have pushed this figure higher. how rapidly) global CO2
theory of Earth’s thermal regulation. For example, the 10 warmest years emissions diminish. Scientists
John Tyndall was the first to on record have occurred since 1998. predict that, at the current
prove Fourier’s greenhouse effect rate, this increase could
hypothesis. His experiments Fueling a warming world range from 0.5°F to 8°F
demonstrated how, when Earth By 1904, Arrhenius had become (0.3°– 4.6°C) by 2100, with
cools down at night—by releasing concerned about the dramatic the greatest warming likely
the heat absorbed from the Sun increase of CO2 due to human to occur in the Arctic regions.
during the day—atmospheric actions—primarily through burning
gases, especially water vapor, fossil fuels, such as coal and oil. He
absorb the heat (radiation) and correctly predicted the influence
cause a greenhouse effect. This that CO2 emissions would have on
keeps Earth’s temperature at an global temperatures, but eventually
came to the conclusion that an
increase in global temperatures
could have a beneficial effect on
plant growth and food production.
The burning of fossil fuels has,
in fact, increased CO2 levels more
If the planet were a patient, quickly than Arrhenius expected,
we would have treated although the planet has warmed
her long ago. less than he predicted. Scientists
Prince Charles understand now that global
warming is having damaging The Perito Moreno glacier in
effects on people and on the Patagonia is one of the few glaciers
that is still growing. The majority
environment, and will continue to are slowly melting, causing sea
do so as long as long as emissions levels to rise worldwide.
continue to increase. ■
204
LIVING MATTER IS
THE MOST POWERFUL
GEOLOGICAL
THE BIOSPHERE
FORCE
E
arth has four interacting and water-based environment, and
IN CONTEXT subsystems: the lithosphere, reaches into extreme habitats, such
Earth’s rigid, rocky outer as the intensely hot mineral-rich
KEY FIGURE
shell; the hydrosphere, which waters around hydrothermal vents.
Vladimir Vernadsky
comprises all water on the planet’s It is often divided into “biomes”—
(1863–1945)
surface; the atmosphere, formed by common major habitats, such as
BEFORE layers of surrounding gases; and deserts, grasslands, oceans, tundra,
1785 Scottish geologist James the biosphere—anywhere that and tropical rain forests.
Hutton proposes that in order supports life, from the ocean depths
to understand Earth, all of its to the highest mountaintops. Earth the superorganism
interactions should be studied. The biosphere’s origins are Ideas about the biosphere began
ancient: fossils of tiny single-celled to emerge in the 18th century,
1875 Austrian geologist microorganisms that date back when the Scottish geologist James
Eduard Suess first uses the 4.28 billion years suggest that it Hutton described Earth as a
term “biosphere” to describe is almost as old as Earth itself. The superorganism—a single living
“the place on Earth’s surface biosphere extends into every land- entity. A century later, Eduard
where life dwells.” Suess introduced the concept of the
biosphere in Das Antlitz der Erde
AFTER (The Face of the Earth). Suess
1928 In Methodology of explained that life is limited to a
Systematics, Russian zoologist zone at Earth’s surface and that
Vladimir Beklemishe warns plants are a good example of the
that humanity’s future is Man is becoming a interactions between the biosphere
irrevocably linked to the more and more powerful and other zones—they grow in the
preservation of the biosphere. geological force, and the soil of the lithosphere, but their
change of his position on leaves breathe in the atmosphere.
1974 British scientist James the planet coincided In The Biosphere (1926), Russian
Lovelock and American with this process. geochemist Vladimir Vernadsky,
biologist Lynn Margulis Vladimir Vernadsky who had met Suess in 1911, defined
first publish their Gaia the concept in much more detail,
hypothesis—the idea of outlining his view of life as a major
Earth as a living entity. geological force. Vernadsky was
one of the first to recognize that
atmospheric oxygen, nitrogen,

See also: The ecosystem 134–137 ■ Biodiversity and ecosystem function 156–157
■ A holistic view of Earth 210–211 ■ The Gaia hypothesis 214–217
Vladimir Vernadsky
Born in 1863, Vladimir
Vernadsky graduated from
St. Petersburg State University
aged 22, and did postgraduate
work in Italy and Germany,
where he studied the optical,
elastic, magnetic, thermal,
Over billions of years layers of oxygen increased, more complex and electrical properties of
cyanobacteria have fossilized to form life forms evolved that would shape crystals. After the revolution
stromatolites—mounds of sedimentary Earth in different ways, eroding in Russia in February 1917,
rock, as seen here at Hamelin Pool, Vernadsky became assistant
Shark Bay, Western Australia.
and remolding its surface, and
Minister of Education in the
changing its chemical composition.
provisional government. The
Gradually, elements of the following year, he founded the
and carbon dioxide result from biosphere became part of the Ukrainian Academy of Science
biological processes, such as the lithosphere. Over millennia, dead in Kiev. Although his book
respiration of plants and animals. corals created reefs in shallow The Biosphere was not taken
He argued that living organisms tropical oceans. Similarly, the calcite seriously by scientists outside
reshape the planet as surely as skeletons of trillions of marine Russia for many years, it later
physical forces, such as waves, organisms fell to the ocean floor, became one of the founding
wind, and rain. He also introduced fossilized, and formed limestone. ■ documents of Gaia theory.
the idea of three stages of Earth’s In the 1930s, Vernadsky
development: first, the birth of the advocated the use of nuclear
planet with the geosphere, in which power, and played an advisory
only inanimate matter existed; role in the development of the
secondly, the emergence of life in Soviet atomic bomb project.
He died in 1945.
the biosphere; and finally the epoch
in which human activity changed I look forward with
the planet forever—the noosphere. great optimism. Key works
We live in a transition
to the noosphere. 1924 Geochemistry
Sphere interactions 1926 The Biosphere
Scientists believe the biosphere Vladimir Vernadsky 1943 “The Biosphere and
has constantly changed. Oxygen the Noosphere”
levels in the atmosphere began 1944 “Problems of
to rise at least 2.7 billion years Biochemistry”
ago, as microorganisms called
cyanobacteria multiplied. As
206
IN CONTEXT
THE SYSTEM
KEY FIGURES
Frederic Clements
(1874–1945),
OF NATURE
Victor Shelford (1877–1968)
BEFORE
1793 Alexander von Humboldt
coins the word “association”
BIOMES to sum up the mix of plant
types that occurs in a
particular habitat.
1866 Ernst Haeckel poses
the idea of the biotope, the
living space for a range of
plants and animals.
AFTER
1966 Leslie Holdridge
champions the idea of life
zones based on the biological
effects of temperature and
rainfall variations.
1973 German–Russian
botanist Heinrich Walter
creates a biome system that
considers seasonal variations.
D
ifferent parts of the world
have varying patterns of
plant and animal life, but
there are usually similarities over
vast areas. These are called biomes,
and each one is a large geographical
region with its own distinctive
plant and animal community and
ecosystem. The idea of the biome
was first popularized by plant
ecologist Frederic Clements and
zoologist Victor Shelford in the US,
in their key book Bioecology (1939),
although its origins date back earlier.
The biome concept took shape
as ideas on plant succession and
community ecology developed.
Clements identified “formations,”
large plant communities, which led

See also: The distribution of species over space and time 162–163 ■ Climax
community 172–173 ■ Open community theory 174–175 ■ Biogeography 200–201
to his idea of climax communities in

1916. The same year, Clements used The geographic spread
the term “biome” to describe biotic of plants is determined
communities—all the interacting mainly by climate.
organisms within a specific habitat.
Like-minded thinkers
Clements was not the only one
thinking along these lines. Zoologist Different plants flourish
Victor Shelford was working toward in each climatic region.
the same idea. The pair began to Threatened coral
meet over the next 20 years, while
reef biomes
pursuing their own research, to see
how they could combine the worlds Coral reefs are such bountiful
of plants and animals. Clements The major types of plants habitats that they are often
studied plant biomes in Colorado in each region match seen as the tropical rain
with his wife, the eminent botanist precipitation forests of the sea. They
Edith Clements. Meanwhile, Shelford and temperature support a quarter of all marine
compiled the Naturalist’s Guide to patterns closely. species and provide livelihoods
the Americas (1926)—the first major for half a billion people. Yet
geographical summary of wildlife in they now face catastrophe.
the Americas, in which he talked Half of all reefs have been lost
about “biota.” This book laid much in the last 30 years, and some
of the foundation for later findings. experts estimate that 90
The major plant percent will be gone over the
Ways of looking at interactions types can be used next 30 years. The main global
in ecological communities took a to divide the world threats are ocean acidification
major step forward when British ❯❯ into broad natural and global warming. As seas
zones called warm, stressed corals expel
The Mongolian steppe belongs to the biomes, which the algae they rely on for food.
same grassland biome as the prairies reflect variations They stop growing, lose their
in North America. Despite being on colour, and often die in what is
in climate. called a coral bleaching event.
separate continents, they are linked
by their climate, animals, and plants. Such events are becoming
ever more frequent. There are
local threats, too, including
overfishing, both for the table
and for aquariums. Even more
seriously, to catch fish for
aquariums, sodium cyanide is
often squirted into the water
to temporarily immobilize the
fish, and this kills corals. More
brutally, fish for the table are
often caught by throwing
dynamite into the water. This
kills fish, making them easy to
scoop up in vast numbers, but
it also blasts coral reefs apart.
208 BIOMES
botanist Arthur Tansley introduced example, in every continent, but rainforest, grasslands, and deserts,
the term “ecosystem” in 1935. most tree species appear only in but there is no agreed definition
When Clements and Shelford one continent. So, the range of trees and there are marked variations.
published the results of their within the Amazonian forests is
collaboration in 1939, they were not completely different from the range The climate factor
making a sudden breakthrough— of trees in the forests of Indonesia. The one common factor in all biome
rather it was a consolidation of Yet both areas are identifiable as classifications has been climate,
ideas that had been taking shape tropical forest, because the trees although other “abiotic” factors can
over a long time. have features in common. also play a part. Climate determines
The collaboration between Since Bioecology first appeared, the form of plant growth best suited
botany and zoology was crucial. there have been countless attempts to a region, and plants that grow
Only by looking at the totality of to define what a biome is, and many in a certain way are restricted
the natural world with its dynamic different ways of classifying them. to particular climates. The leaves
interactions could scientists hope Biomes provide a simple way of deciduous trees are broad, with
to get a full picture, and Clements of understanding global vegetation a large surface for light absorption,
defined a biome as “an organic unit patterns, but when looked at closely but little resistance to drying out
comprising all the species of plants they present a crude way of grouping or frost. Conifer tree needles, on the
and animals at home in a particular ecosystems. There is no single other hand, are narrow and can
habitat.” Even so, biomes have accepted classification system, and survive the harshest frosts. Desert
come to be defined principally the only division everyone seems to shrubs often have very thin leaves,
by vegetation type. agree on is that between terrestrial or no leaves at all, to resist drying
The most important feature of (land-based) and aquatic (water- out. Biogeographers acknowledge
biomes is that they link vegetation based) biomes. Many of the same climate’s key role when they talk
and plant communities across the biomes crop up in most systems, about “tropical” rainforests and
world. There are tropical forests, for such as the polar biome, tundra, “temperate” grasslands.
Terrestrial biomes of the world
Tropical forest
Temperate forest
Mountains
Grasslands This map shows six biomes across the globe. Each area
has distinct flora, as major plant types vary from one climatic
Desert
region to another. Ocean and freshwater biomes are not
Polar regions displayed here, but are equally important to the biosphere.

Ecozones
Biomes are about identifying
the similar forms that life
takes in response to particular
regional conditions such as
climate, soil, and topography.
However, there are other
methods of dividing the world
in ecological terms. In 1973,
Hungarian biologist Miklos
Udvardy came up with the
concept of biogeographic
realms; this system was then
further developed in a scheme
by the World Wildlife Fund.
The BBC later replaced the
Tropical rainforest is the hottest and of the pyramid represent three term “biogeographic realm”
wettest biome and covers 7 per cent axes: rain, temperature, and with “ecozone.” Biogeographic
of Earth’s surface. One of the oldest realms divide the whole planet
evapotranspiration (which depends
biomes, it also contains far more animal according to the evolutionary
and plant species than any other biome.
on both rain and temperature). Using history of plants and animals.
these axes, he could plot hexagons The ways in which continents
showing regions that also reflect have split apart and drifted
Very few species have identical humidity, latitude, and altitude. means that species have
climate requirements. Even among American plant ecologist Robert evolved variously in different
varieties of the same plants, there Whittaker devised a much simpler parts of the world. Ecozones
are variations. The sugar maple graph, with average temperature are therefore based on
of eastern North America, for on one axis and annual rainfall on identifying this diversification.
example, is slightly more tolerant the other. With these two variables Australasia, for example, is
of winter cold than its cousin the plotted against each other, he was a single ecozone, because
silver maple. Although the areas able to divide the graph into nine marsupials evolved there in
where both trees grow overlap, the biomes—from tropical rain forest isolation from other mammals
sugar maple can be seen far over the (the hottest and wettest) through in the rest of the world.
Canadian border, whereas the silver to tundra (the coldest and driest).
maple flourishes as far south as Underpinning all these systems
Texas. Since biomes give only an is the idea of convergent evolution,
approximate picture of plant and which argues that species develop
animal distribution, ecologists are similar traits as they adapt to similar
constantly devising new systems environments. Insects, birds, bats,
of classification. and pterosaurs all developed wings
independently to occupy air space.
Rain, heat, and evolution Different biomes are therefore
One of the most widely recognized assumed to develop corresponding
systems of classification is the life life forms in response to similar
zones system devised by American environmental conditions. However,
botanist Leslie Holdridge in 1947, in recent decades, it has been noted
and updated in 1967. His system is that species can evolve differently
based on the assumption that two in the same biome and also that The short-beaked echidna
key factors, rain and heat, determine different stable biomes can develop is one of the most widespread
native mammals in the Australasian
vegetation type in each region. He in an identical climate. While central ecozone. They live in a range of
created a graphic depiction of 38 life to understanding life, biomes remain habitats from desert to rain forest.
zones in a pyramid. The three sides a complex and elusive concept. ■
210
WE TAKE NATURE’S
SERVICES FOR GRANTED
BECAUSE WE DON’T PAY
FOR THEM
A HOLISTIC VIEW OF EARTH
IN CONTEXT
KEY FIGURE
Eugene Odum (1913–2002)
BEFORE
1905 In Research Methods in
Ecology, American botanist
Frederic Clements writes
about plant communities and
how they change over time.
1935 Arthur Tansley, a British
botanist, proposes the term
“ecosystem” to describe a
community of plants, animals,
soil minerals, water, and air.
AFTER
1954 Eugene and Howard
Odum’s study of the coral
T
Eniwetok Atoll in the Pacific he American ecologist Salt marshes, such as these on the
Ocean applies the principles Eugene Odum was not coast near Porthmadog, North Wales,
of holistic ecology. the first scientist to write form their own ecosystem, with the
seawater and its nutrients providing
about ecology, but in the 1950s he
1974 British environmentalist a unique habitat for wildlife.
proposed that it deserved to be a
James Lovelock and American discipline in its own right. Until
biologist Lynn Margulis first then, ecology was viewed as a could never lead to a full knowledge
publish their Gaia hypothesis. relatively insignificant subdivision of the living world. He argued that
It states that Earth is a self- of the biological sciences—the poor it was more important to study the
regulating system that relation of biology, zoology, and places and roles that the species
maintains the conditions botany. However, Odum believed held in their community, rather
necessary for life on our planet. passionately that studying plant than simply finding out more
and animal species in isolation about what they were. Odum’s

See also: The ecosystem 134–137 ■ Macroecology 185 ■ The peaceful
coexistence of humankind and nature 297 ■ The Green Movement 308–309
Earth Day
After witnessing a horrific
oil spill in Santa Barbara,
new approach to the subject—
California, in 1969, US Senator
first set out in his 1953 book Gaylord Nelson decided to
The Fundamentals of Ecology— focus on growing worries
revolutionized the purpose and about pollution during a
influence of ecological research. national “teach-in” on the
…ecology has been environment. He could not
The “new ecology” badly presented and have envisaged the size of the
The holistic view of Earth involves has been broken into movement he would inspire.
studying the systems of organisms too many antagonistic On April 22, 1970, 20 million
as a whole. As Odum explained, subdivisions. Americans took part in the
one organism, or any one group of Eugene Odum first Earth Day, with rallies,
organisms, cannot be understood marches, and lectures taking
without studying the ecosystem in place nationwide. Such was
which it lives. The holistic approach the effect of the protests that
later that year the Clean Air,
examines all the roles played by
Clean Water, and Endangered
each member of an ecosystem,
Species Acts became law, and
and how that system interacts with the Environmental Protection
others. Climate, geology, water and the whole animal, or the ecosystem Agency was established in the
mineral input, and human activity in which the animal lives—are able US that December. Earth Day
all affect—and are affected by—a to self-regulate to provide stability. became a global phenomenon,
multitude of living communities. with 200 million people
Odum was writing in the 1950s Integrated investigation participating in 141 countries
and ’60s, when there was a growing A holistic study of a lake ecosystem in 1990—and built momentum
awareness of the environmental would involve looking at all the for the 1992 UN Earth Summit
destruction wrought by humanity. inputs into the lake and its margins in Rio de Janeiro. Earth Day
The role of people was a crucial part as well as all the outputs, including celebrations are held every
of “systems ecology,” as he called energy, water, minerals, and April, with a different theme
his idea. Odum wanted humans to nutrients. It would also consider each time. In 2018, the focus
be sympathetic allies with the any human inputs. The study was on ending global pollution
by plastics.
natural world—collaborators rather would examine the roles played
than manipulators—and his views by both producer organisms, such
of an all-embracing ecology did as plants and algae, and consumers
much to inspire the first Earth Day, such as herbivores and carnivores.
which was celebrated in 1970. The holistic approach also examines
The holistic concept of Odum’s changes over time, in which
“new ecology” deals with Earth developments that benefit some
as a whole, bringing together organisms in the short-term might
physics, chemistry, botany, zoology, lead to a lack of diversity in the
geology, and meteorology. The future. For example, although trout
fundamental assumptions of thrive in warmish, alkaline waters,
ecology are that the ecosystem if those waters become too warm
is the basic unit of nature, that or acidic due to ecological change,
biological diversity increases the the fish can no longer breed. The first Earth Day on April 22,
ability of ecosystems to survive, Odum’s holistic approach leaves 1970, saw crowds such as this
and that the whole is greater than a legacy of a far more detailed one in Philadelphia, Pennsylvania,
gather across the US to protest
the sum of its parts. Systems in the appreciation of what is happening against pollution and the use
natural world—whether they are in an ecosystem than a series of of pesticides.
groups of cells in an animal’s body, individual species studies. ■
212
PLATE TECTONICS
IS NOT ALL HAVOC
AND DESTRUCTION
MOVING CONTINENTS AND EVOLUTION
T
he surface of Earth is
IN CONTEXT constantly moving, very
KEY FIGURE slowly, and has been doing
so for more than three billion years.
Alfred Wegener (1880–1930)
The lithosphere (Earth’s crust and
BEFORE upper mantle) is divided into seven
1596 Abraham Ortelius, a large sections and many smaller
Dutch scholar, is one of several ones, called tectonic plates. Where
geographers who observe that plates meet, the type of movement
the two sides of the Atlantic determines the nature of the This fossilized head of the extinct
boundary. Where plates push reptile Cynognathus crateronotus was
Ocean seem to “fit” each other. found in southern Africa. The same
against each other, new mountains
fossils occur in South America: evidence
AFTER are created. If plates pull apart, new that the two continents were once one.
1929 British geologist crust forms on the ocean floor.
Arthur Holmes proposes The first inkling that the
that convection in Earth’s continents may not have always animals or plants concerned would
mantle drives continental drift. been in their current positions came have been unable to cross the ocean
in the late 16th century. European divide. These include Cynognathus
1943 George Gaylord Simpson explorers sailing to the Americas crateronotus, a mammal-like reptile
dismisses fossil evidence for saw from their newly created maps that lived over 200 million years ago
continental drift and argues that the coastlines on each side in southern Africa and eastern South
for “stable continents.” of the Atlantic Ocean mirrored America. Glossopteris, a genus of
each other. Later, geologists found woody trees, grew in South America,
1962 American geologist
strong structural and geological South Africa, Australia, India, and
Harry Hess explains how the
similarities between the Caledonian- Antarctica, but nowhere else, around
seafloor spreads, by molten
era mountains of Northern Europe 300 million years ago.
magma rising from below. and the Appalachian Mountains of To German geophysicist Alfred
2015 A group of Australian North America. Wegener, such fossil patterns
scientists propose that periods indicated that these continents had
of rapid evolution in the oceans Lookalike fossils once been joined together. In 1915,
were triggered by collisions There are various examples of fossil he published his theory that all the
between tectonic plates. finds straddling different continents continents were once a single land
that can only be explained by mass, “Pangaea,” which has since
continental movement—since the broken up and drifted apart.

See also: Island biogeography 144–149 ■ The distribution of species over space and time 162–163 ■ Macroecology 185
■ Metapopulations 186–187 ■ Biogeography 200–201
Three types of plate boundary

Plates slide
Plates move apart Plates collide alongside each other
Tectonic plates can move in three

different ways, forming different types
of boundary. When plates diverge, new
oceanic crust is formed. When they
converge, new mountains form. When
plates slide past each other, the rift is
known as a transform fault. Divergent Convergent Transform
Wegener’s theory was not well plates is driven by convection

received at first. In 1943, George currents carrying heat from deep
Gaylord Simpson, one of the most inside the planet to the surface.
influential paleontologists in the Once Wegener’s theory was
US, criticized the theory. He argued accepted, the fossil evidence made
that the fossil record could be much better sense. Continental drift The forces which
explained by static continents linked has had a profound influence on how displace continents
and unlinked by periodic flooding. species have evolved. For example, are the same
if a continent splits apart, the two as those which
Evidence and evolution separated populations of a species produce great
Despite early doubts, evidence for can start to evolve in completely fold-mountain ranges.
the plate tectonics theory grew. A different directions. On the other Alfred Wegener
series of discoveries established that hand, if two continents collide, or a
the seafloor was spreading and that bridge of land forms between them,
new oceanic crust was constantly different species begin to mix and
being created. We now understand compete, and some may become
that the movement of the tectonic extinct as a result. ■
Marsupials in America and Australia

Marsupials are nonplacental thought that they traveled via a
mammals whose young complete belt of vegetation straddling the
their gestation feeding from their three areas, which were once all
mother’s teats, typically in a part of the southern landmass
pouch on the belly. Now found called Gondwana.
only in the Americas (mainly By 55 million years ago,
South and Central) and Australia, the continents had separated,
they are thought to have evolved and marsupial species began
in North America 100 million to evolve differently. The only
years ago. They spread to South known Antarctic marsupial
America and diversified into many fossils, found on Seymour Island
different species. in 40-million-year-old rocks,
Marsupials are strongly identified Several groups later moved resemble South American
with Australia, yet they evolved in into what is now Antarctica and marsupials of the same period,
America and are still also found there. on into southern Australia. It is but not those of Australia.
214
IN CONTEXT
LIFE CHANGES
KEY FIGURE
James Lovelock (1919–)
EARTH TO ITS
BEFORE
1935 British botanist Arthur
Tansley uses “ecosystem” to
describe an interdependent
OWN PURPOSES
community of biological and
nonbiological components.
1953 In Fundamentals of
Ecology, American ecologist
THE GAIA HYPOTHESIS Eugene Odum describes
Earth as a collection of
interlocking ecosystems.
AFTER
1985 In the US, the first
conference on the Gaia
hypothesis is held, entitled,
“Is the Earth a Living
Organism?”
2004 James Lovelock voices
his support for nuclear power
over renewable energy.
I
n 1979, British scientist James
Lovelock’s book Gaia: A New
Look at Life on Earth presented
his Gaia hypothesis to a general
readership. In essence, Lovelock
claimed that Earth is a single, self-
regulating system, in which living
and nonliving elements combine
to promote life. The book quickly
became a bestseller, and caught
the imagination of the growing
Green movement, offering a fresh
approach to environmentalism.
What Lovelock proposed was
not without precedent. In the 1920s,
Vladimir Vernadsky, a Russian
scientist, had developed the idea
of the biosphere, the zone of Earth
that holds all living organisms, and
suggested that it could be seen as
a single entity in which organic and

See also: The ecosystem 134–137 ■ Evolutionarily stable state 154–155
■ The biosphere 204–205 ■ A holistic view of Earth 210–211
pedosphere, the surface layer of the

Earth; the hydrosphere, the bodies of
water on the Earth’s surface; and the
atmosphere, the gases surrounding
the Earth. These spheres and their
Evolution is a tightly coupled complex interactions maintain
dance, with life and the Earth in “homeostasis.” This
material environment as concept is borrowed from physiology,
partners. From the dance which describes the stable internal
emerges the entity Gaia. conditions, such as temperature
James Lovelock and chemical composition, that James Lovelock
allow organisms to function
optimally. They are controlled by Inspired by writers such as
self-regulating mechanisms Jules Verne and H.G. Wells,
that react to change in those James Lovelock, born in 1919,
conditions. Lovelock’s use of the was fascinated by science and
word homeostasis reinforced invention from an early age.
inorganic elements interact. The the implication that Earth, He graduated in chemistry
British botanist Arthur Tansley or Gaia, is a living entity. from Manchester University
then took this idea further in the in 1941. Lovelock was a
1930s, with his concept of an Keeping the balance conscientious objector during
“ecosystem” that regulates itself The hint of mysticism in the Gaia World War II and worked for
the National Institute for
into a state of equilibrium. principle chimed with the “New
Medical Research in London.
Tansley’s theory was at the Age” thinking of the time. This
In 1948, he received his Ph.D.
heart of Lovelock’s hypothesis: helped popularize the idea, but in medicine, and then spent
that all living organisms and their it also led to a negative reception time in the US on a Rockefeller
environment form one complex from the scientific establishment. fellowship. After returning
super-ecosystem that regulates and However, behind the Earth to Britain in 1955, he turned
balances conditions to sustain life “goddess” metaphor was a serious his attention to inventions,
on Earth. The idea first occurred to science-based hypothesis that ❯❯ notably the electron-capture
Lovelock in the late 1960s, but it detector (ECD), which detects
was after discussing it with US trace atoms in a gas sample.
microbiologist Lynn Margulis that In the 1960s and 1970s, he
it began to take shape. Together, held visiting professorships in
they presented the hypothesis in a Houston, Texas, and Reading,
paper in 1974, giving it a name England, during which time
suggested by the writer William he developed the Gaia
hypothesis. In 2003, Lovelock
Golding—Gaia, after the ancient
was made a Companion of
Greek Earth goddess. Lovelock and Honour by Queen Elizabeth II.
Margulis portrayed Earth as a
living entity, composed of the
biosphere, living organisms; the Key works
1988 The Ages of Gaia

A stone relief shows Gaia, the 1991 Gaia: The Practical
Greek goddess of Earth. The Science of Planetary Medicine
nonscientific name chosen by Lovelock 2009 The Vanishing Face of
for his hypothesis initially hindered Gaia: A Final Warning
its acceptance by many scientists.
216 THE GAIA HYPOTHESIS
Daisyworld
At first, scientists criticized
the Gaia hypothesis for its
supposed implication that the
ecosystems in the biosphere
could collectively influence
Earth’s environment. So to
enhance the plausibility of the
Gaia theory, in 1983 James
Lovelock and fellow British
scientist Andrew Watson
produced “Daisyworld,” a
simple explanatory model.
Daisyworld is a barren
planet, orbiting a sun. As the
intensity of the sun’s rays
increases, black daisies start
to grow. They absorb heat and
warm the planet’s surface to
the point where white daisies
can thrive. They, in turn,
reflect the sun’s energy, so
cooling the ground. The two In the Gaia hypothesis, Earth, the different state of equilibrium. Such
kinds of daisy reach a point only known planet to support life, is a tipping point, argued Lovelock,
itself a “superorganism,” where the sea, occurred about 2.5 billion years
of equilibrium, whereby they
land, and atmosphere work together to
regulate the temperature of maintain the right living conditions.
ago, at the end of the Archean Eon,
the planet. When the sun’s when oxygen first appeared on
heat increases further, the Earth. At this time, Earth was a
white daisies, able to reflect salinity in its environment. When hot, acidic place in which methane-
the sunlight and stay cool, these are constant, Earth is in producing bacteria were the only
replace the black daisies. a stable state of homeostasis, but life that thrived. Bacteria capable of
Finally, the sun heats up so if the balance is disturbed, the photosynthesis then evolved, which
much that even the white planet encourages the organisms created an atmosphere that was
daisies can no longer survive.
that will restore the equilibrium,
while being hostile to those that
the interactions of living organisms reinforce the disturbance. The
and their physical surroundings— organic components of the Earth
including the cycles of oxygen, system do not simply react to
carbon, nitrogen, and sulfur— changes in their environment, but
form a dynamic system that control and regulate them. If there were
stabilizes the environment. These feedback mechanisms a nuclear war,
According to Lovelock, Gaia operate in a complex global network and humanity were
is controlled by the action of of interconnected natural cycles, to wiped out, Earth
“feedback loops,” which are maintain the optimum conditions would breathe
the checks and balances that for the organisms within them. a sigh of relief.
compensate for disturbances in They can resist change, but only James Lovelock
the system, bringing it back into to a certain extent. A big enough
equilibrium. To function well, life disturbance can push the system
on Earth depends on a particular to a “tipping point,” where, with the
balance of variables such as water, balance of its components altered,
temperature, oxygen, acidity, and it is likely to settle into a very

conducive to more complex forms An algal feedback loop
of life. Eventually, the equilibrium
conditions that exist on Earth
today were established. Lots of clouds mean More DMS
heat from the Sun means
Saving the planet is reflected back more cloud.
As Lovelock elaborated on the into space.
theme, the scientific establishment
gradually began to accept the Gaia
hypothesis. In the 1980s, a series
of “Gaia conferences” attracted
scientists from many different
disciplines, willing to explore the
mechanisms involved in regulating Cooler temperatures
Warmer temperatures
Earth’s environment to achieve mean lower algal
mean higher algal growth
homeostasis. Later, more attention growth and
and more DMS.
was devoted to looking at the less DMS gas.
implications of the hypothesis
in the face of climate change.
Human activity had been shown
to disturb Gaia’s system, but the
issue was now whether its
regulatory mechanisms could Less DMS Fewer clouds mean
withstand further pressure—or means more heat from the Sun
whether Earth was facing another fewer clouds. reaches Earth.
irreversible tipping point.
Environmentalists, who had
been among the first to embrace
In Gaia theory, feedback loops keep Earth in
Gaia, reacted with dismay to the balance. One example is the effect that sea algae called
theory that the human species may coccolithophores have on keeping the planet’s climate in
precipitate a catastrophic change check. When the algae die, they release a gas, dimethyl
in Earth’s equilibrium. The rallying sulphide (DMS), that helps to create clouds.
cry of Green activists became
“Save the planet!” but this was at
odds with the fundamental idea of
Gaia. Although the destruction
of natural habitats, the excessive
burning of fossil fuels, the depletion
of biodiversity, and other human-
made threats were likely to have
severe consequences for many
species—including humans—
the planet, according to the Gaia
hypothesis, will survive and find
a new equilibrium. ■
Nuclear power stations produce

plentiful “clean” energy, but also toxic
waste. James Lovelock believes Earth
is able to absorb and overcome the
waste’s radioactive effects.
65 MILLION
YEARS AGO SOMETHING
KILLED HALF
OF ALL THE LIFE ON
THE EARTH
MASS EXTINCTIONS
220 MASS EXTINCTIONS
IN CONTEXT
KEY FIGURE
Luis Alvarez (1911–88)
BEFORE
1953 American geologists
Allan O. Kelly and Frank
Dachille suggest in their book
Target: Earth that a meteor
impact may have been
responsible for the extinction
of the dinosaurs.
AFTER
1991 The Chicxulub Crater
in the north of the Yucatan
Peninsula in southeastern
T
Mexico is proposed as the site
of a massive comet or meteor here have been five periods The meteor that hit Earth at the end
in Earth’s history when of the Cretaceous period was traveling
impact at the end of the at 40,000 mph (64,000 kph). Its power
Cretaceous period. abnormally large numbers
was a billion times greater than the
of multicellular organisms have Hiroshima atomic bomb.
2010 An international panel died off in a relatively short time.
of scientists agrees that the These mass extinctions are defined
Chicxulub impact led to by the loss of multicellular animals five families of marine animals every
the Cretaceous–Paleogene and plants because their fossils are million years. This is far exceeded
mass extinction, around far easier to detect than those of during mass extinctions, which
65 million years ago. single-celled organisms. always mark the boundary between
The general (“background”) rate two geological periods. Scientists
of extinction is between one and five do not understand all the factors
species a year. Fossil records show, responsible for these events, though
for example, the extinction of two to they are agreed on some. Increased
Mass extinction events from

499 million years ago to the present
Late Devonian Triassic
A rapid drop in sea level Climate change or
is one of a number of an asteroid hit are
possible causes for the potential causes for
loss of 70–80 percent the extinction of round HOLOCENE
(OR ANTHROPOCENE) PERIOD
of animal species. 75 percent of species. 100,000 YEARS AGO–PRESENT
ORDOVICIAN SILURIAN DEVONIAN CARBONIFEROUS PERMIAN TRIASSIC JURASSIC CRETACEOUS (K) PALAEOGENE NEOGENE
485–444 444–419 419–359 359–299 299–252 252–201 201–145 145–66 66–23 23–03
Ordovician Permian Cretaceous

Global cooling Huge volcanic A meteor strike and
leads to the activity helps to volcanic activity drive
extinction of wipe out 96 percent up to 80 percent of
85 percent of of all marine species. animals, including
marine life. most dinosaurs,
to extinction.

See also: Ancient ice ages 198–199 ■ Moving continents and evolution 212–213 ■
The Gaia hypothesis 214–217 ■ Ocean acidification 281
on Earth but common in asteroids.

The discovery led to the Alvarez
Hypothesis, which proposed that
the extinction at the end of the
Cretaceous period was caused
All geologic history is full by a catastrophic meteor strike.
of the beginnings and the The location of the impact was still
ends of species–of their a mystery, until 11 years later, when
first and last days. a massive crater 106 miles (170 km)
Hugh Miller across on Mexico’s Yucatan
Scottish geologist Peninsula was found to date from Luis Alvarez
the time of the extinction.
The scientific consensus is that Considered one of the greatest
a massive comet or asteroid struck physicists of the 20th century,
Earth, producing a blast of radiation Luis Alvarez was born in
and a destructive megatsunami San Francisco in 1911. He
more than 328 ft (100 m) high. The graduated from the University
volcanic activity, changes in the radiation would have killed animals of Chicago in 1936 and went
composition of the atmosphere and nearby, and the megatsunami on to work at the Radiation
oceans, climate change, sea level would have obliterated coastal Laboratory in the University
rises and falls, tectonic movement of regions around the Gulf of Mexico. of California, Berkeley. There
the continents, and meteor impacts The main damage, however, would he helped develop nuclear
reactors and, during World
are all likely causes. Some scientists be more gradual. A vast cloud of
War II, nuclear weapons. He
suggest we have now entered a soot and dust would have spread
witnessed the atomic bombing
sixth mass extinction, this time the through the atmosphere, blocking of Hiroshima and helped build
result of human activity. out sunlight for several years. Plants a plutonium bomb.
died because they could no longer After the war, Alvarez
End of the dinosaurs photosynthesize, and algae in coral developed the liquid hydrogen
The mass extinction that scientists reefs also succumbed, disrupting bubble chamber, used to
understand best is also the most food chains worldwide. The ❯❯ discover new subatomic
recent, around 65 million years ago. particles. For this, in 1968 he
Geologists refer to it as the K-Pg was awarded the Nobel Prize
extinction event because it occurred for Physics. Later he provided
at the end of the Cretaceous and the calculations to back up the
start of the Palaeogene periods. Alvarez Hypothesis of mass
Although an extraterrestrial origin extinction caused by a meteor
was first suggested for the event in We have very strong physical strike. He died in 1988.
the 1950s, this was not taken and chemical evidence for
seriously until two discoveries, in a large impact … the Key works
Europe and North America. extinction coincides with
In 1980 a team of scientists the impact to a precision 1980 “Extraterrestrial Cause
working in Italy, including physicist of a centimetre or better. for the Cretaceous–Tertiary
Luis Alvarez and his geologist son, Walter Alvarez Extinction,” Science
1985 “The Hydrogen Bubble
Walter Alvarez, discovered a clay
Chamber and the Strange
layer between Cretaceous and Resonances”
Paleogene deposits. Examination 1987 Alvarez: Adventures
of the clay revealed that it contained of a Physicist
the mineral iridium, which is rare
222 MASS EXTINCTIONS

Although many flying dinosaurs
survived the K-Pg mass extinction
at the end of the Cretaceous period,
all pterosaurs perished, ending their
162-million-year stay on Earth.
dramatically, toward the end

of the Ordovician period, around
444 million years ago. At this time,
most organisms on Earth lived in
the oceans. As the supercontinent
Gondwana drifted slowly over the
South Pole, a giant ice cap formed,
lowering global temperatures.
Much of the planet’s water became
“locked up” as ice, depressing sea
levels and reducing the area of
Earth’s surface covered by ocean.
As a result, marine organisms
living in shallow continental-shelf
water suffered particularly high
rates of extinction. In at least two
peak die-off periods, separated
by hundreds of thousands of years,
nearly 85 percent of marine species
died out, including brachiopods,
bryozoans, trilobites, graptolites,
and echinoderms.
impact would have also released with a fast metabolism that
sulfuric acid into the atmosphere, demanded regular meals. Many Slow extinction
which produced acid rain, plant species died because they By the Late Devonian period,
acidifying the oceans and killing could not photosynthesize, leaving around 359 million years ago, the
off marine life. Around the same herbivorous dinosaurs with little continents had been colonized by
time, a huge amount of volcanic vegetation to eat, while predatory
activity flooded 193,000 sq miles species starved for lack of prey. In
(500,000 sq km) of southern India contrast, fungi, which do not depend
with lava, forming the Deccan on photosynthesis, proliferated.
Plateau and further changing the In the oceans, phytoplankton, a
climate and atmosphere. vital food source that also relied on The current
The K-Pg event is best known photosynthesis, died out. Creatures extinction has its
for the extinction of all nonflying that fed on phytoplankton then own novel cause: not
dinosaurs. It was also responsible faced extinction. These included an asteroid or a massive
for the death of nearly all four- cephalopods, such as belemnites
volcanic eruption but
legged animals (tetrapods) that and ammonites, and the marine
weighed more than 55 lb (25 kg). reptiles known as the mosasaurs
“one weedy species.”
An exception were crocodiles, and the sauropterygians.
Elizabeth Kolbert
American journalist
which may have survived because
they are ectotherms (cold-blooded Marine annihilation
animals), able to survive for a long The earliest mass extinction,
time without food. Dinosaurs were and the second-most catastrophic,
endotherms (warm-blooded animals), occurred when our planet cooled

plants and insects, and massive Earth’s history. The eruptions,
organic reefs thrived in the oceans. which lasted nearly 1 million years,
The continents of Euramerica and flooded more than 0.8 million sq
Gondwana were converging into miles (2 million sq km) of ancient
what would become Pangaea— Siberia with basalt lava. The
the last of the supercontinents. Modelled future extinction resulting buildup of greenhouse
In this period, a succession of rates are projected to gases would have transformed the
extinctions—possibly as many be 10,000 times Earth’s atmosphere of Earth, likely resulting
as seven—took place over a longer historical geological in severe global warming and
timescale than any other mass background rate. contributing to species extinction.
extinction event, possibly up Ron Wagler
to 25 million years. American academic Phased losses
The extinctions may have had All life today is descended from
many causes, including reduced the small minority of species
oxygen in the oceans, falling sea that remained at the start of the
levels, atmospheric changes, the Triassic period. During the
draining of water produced by the period’s final 18 million years,
spread of plants, and asteroid ending about 201 million years
impacts. Most organisms lived in known as “The Great Dying,” it ago, at least half of all animal
the oceans, and shallow seas were resulted in the loss of 96 percent species known to be living at that
worst affected, with many reef- of marine species and 70 percent of time were wiped out in two or three
building organisms, brachiopods, land-living vertebrates. Insects extinction phases. Climate change
trilobites, and the last of the suffered the only mass extinction in caused by more basalt eruptions
graptolite species dying off. Around their history, and the last of the and an asteroid impact have been
75 percent of marine species died, trilobites, which had been in cited as causes. In the seas, many
and it would be another 100 million decline for millions of years, reptiles, cephalopods, mollusks,
years before corals re-established disappeared from the fossil record. and reef-building organisms died
themselves on a large scale. Potential causes for the mass out. On land, most of the reptilelike
extinction include asteroid impact archosaurs and many large
“The Great Dying” and oxygen depletion in the amphibians became extinct. The
The most dramatic mass extinction oceans. The extinction also loss of the archosaurs, in particular,
took place at the end of the Permian coincided with one of the biggest opened up ecological niches that
period, 252 million years ago. Also periods of volcanic activity in the dinosaurs would fill. ■
The sixth extinction These losses have been driven by

habitat change, climate change,
Some ecologists have estimated overfishing, overhunting, ocean
the current rate of extinction of acidification, air pollution, and
animals and plants at 100–1,000 the introduction of animals that
times the natural background disrupt food chains. American
rate, with most of the increase ecologist E.O. Wilson, known
due directly or indirectly to as “the father of biodiversity,”
human activities. They argue believes that if the species die-off
that this is evidence the world continues at the present rate, half
is already in the middle of the of all higher life forms will be
Holocene extinction, named for extinct by 2100. Stuart Pimm,
the present geological epoch. a British–American biologist and
Many species of animals and modern extinctions expert, is Sudan, the last male northern
plants have been lost since more cautious, claiming that we white rhinocerous, died in 2018 (two
the start of the Industrial are on the cusp of such an event females remain). Poaching has taken
Revolution in the 18th century. and can still act to stop it. the species to the edge of extinction.
224
BURNING ALL FUEL

RESERVES WILL
INITIATE THE RUNAWAY
GREENHOUSE
ENVIRONMENTAL FEEDBACK LOOPS
A
ll parts of an ecosystem
IN CONTEXT are interdependent. Any
species or habitat change
KEY FIGURE Negative feedback loops will feed back into the system, and
James Hansen (1941–) regulate ecosystems. affect the whole of that system,
BEFORE including the part where it all
1875 In the book Climate and started. In other words, the
Change, Scottish scientist feedback travels around in a loop.
James Croll describes the In some situations, change
climate-warming feedback is kept in check by the loop. For
effect of melting ice. example, if aphids suddenly
multiply, they provide more food
1965 Canadian biologist They damp
for ladybugs, leading to an increase
down change.
Charles Krebs discovers the in the number of ladybugs. But
“fence effect,” showing vole with more ladybugs feeding on the
populations protected from aphids, aphid numbers drop again.
foxes rocketing, then crashing. This is negative feedback and it
helps maintain the status quo.
1969 American planetary In other cases, feedback can
scientist Andrew Ingersoll accelerate change. Shrubs, for
highlights the “runaway example, may begin to take over
greenhouse effect” that caused They keep populations
under control. from grass on newly colonized land,
the planet Venus to heat up. casting their shade over the grass,
depriving it of sunlight, and slowing
AFTER
its growth. The shrubs now have
2018 Ecologists in Alaska
more water and nutrients, so they
predict that the accelerating prosper at the expense of the grass.
release of methane from This is positive feedback and it is
formerly frozen lakes will inherently destabilizing.
increase global warming. Negative Ideas about feedback loops first
feedback loops developed early in the 20th century.
provide stability. They were based on the work of
two mathematicians—Alfred Lotka
(1880–1949) in the US and Italian

See also: Predator–prey equations 44–49 ■ Competitive exclusion principle
52–53 ■ Global warming 202–203 ■ Halting climate change 316–321
Feedback loops and
climate change
In recent years, accelerating
and decelerating warming
trends have brought the idea
of feedback loops to the fore
in climate change science.
In 1988, the climate scientist
James Hansen spoke at a US
congressional hearing of the
rises in global temperature
caused by human activity.
He has since voiced the belief
that the continued burning
of fossil fuels could set in
motion a series of calamitous
positive feedbacks on Earth’s
climate, leading to the
“runaway greenhouse” he
describes in his 2009 book
Storms of My Grandchildren.
Vito Volterra (1860–1940)—who In a healthy ecosystem, a repeating
fluctuation in numbers between prey, One warming feedback
independently devised equations loop is created by the melting
based on the interaction between such as rabbits, and predators, such
as foxes, is an example of a negative of polar ice caps, as newly
predators and prey. Their equations feedback loop balancing the system. exposed land and water
showed that a prey population will absorb the heat that the ice
grow rapidly when the number of once reflected back into the
predators drops, while the predator Positive feedback interferes with atmosphere. The melting of
population will drop when prey a balanced ecosystem. If there Siberian permafrost is another
numbers drop, because the is a surplus of resources, or a lack warming loop. As temperature
predators go hungry. The result is of predators, a population can grow rises melt the permafrost,
a constant cycle of falling and freely. A bigger population leads to huge amounts of methane,
rising predator and prey populations. more births, and so an acceleration a greenhouse gas, could be
of the growth in population. released into the atmosphere,
Balancing the system Equally, positive feedback can accelerating global warming.
The predator–prey cycles identified result in an accelerated contraction
by Lotka and Volterra were focused of a population. If fish stocks
on the interaction between single decline in a lagoon, for instance,
predator and prey species. Since local people may resort to importing
their studies, the theory of feedback canned food. Pollution from the
loops has developed to embrace dumps where the cans are thrown
entire ecosystems. Ecologists now away can seep into the lagoon,
think that negative feedback loops killing the fish—and encouraging
are of central importance for the the locals to import even more
functioning of all ecosystems, of the damaging cans. And yet,
keeping every part of them naturally positive feedback loops can
within the bounds of sustainability. sometimes set off a chain of events Arctic areas such as Greenland
Populations can never swell for long that becomes a “virtuous” circle. have seen a reduction in summer
beyond the carrying capacity of the For example, if shrubs are planted ice of 72 percent since 1980. The
warming of the atmosphere and
rest of the system to support them. in unstable soil, their roots may rising sea levels are part of the
Thus, negative feedback regulates stabilize the soil, allowing both resulting positive feedback loop.
an ecosystem and keeps it stable. the shrubs and soil to thrive. ■
THE HUM
FACTOR
AN
228 INTRODUCTION
During the Industrial The world’s first Charles Keeling Gene Likens begins Chico Mendes
Revolution, London’s national park is starts to record the work to establish lobbies the US
“Great Stink” created in the US year-on-year rise the relationship Congress to stop
prompts legislation at Yellowstone in atmospheric between water funding projects
to curb air and to preserve its carbon dioxide quality and that damage the
water pollution. natural habitat. levels. life forms. Amazon rain forest.
1858 1872 1958 1963 1987
1859 1955 1962 1979
Rabbits are released The term “urban Rachel Carson’s The One Child
in Australia; their sprawl” is used for book Silent Spring Policy is initiated
population explosion the first time, by The exposes the harmful in China to
results in chaos for the Times newspaper in effects of pesticides control the rapid
environment. the UK. on the environment. population growth.
R
aw sewage produced Scottish-American environmentalist many forest-dwelling species died
by millions of Londoners John Muir was one of the first to out before they were “discovered.”
once poured into the identify habitat degradation and Deforestation also contributes
Thames River for decades, until destruction as problems, and in to global climate change. As trees
the stench of the effluent became 1890 he won protection for the photosynthesize, they absorb
so bad that in 1858 action was Yosemite Valley in California. carbon dioxide and release oxygen.
demanded. When a new system However, despite a steady increase However, less forest means that
of sewers, pumping stations, and in protected natural environments, more CO2 stays in the atmosphere,
treatment works revolutionized the in the 20th century, the destructive fueling the greenhouse effect and
city’s sanitation, deaths and illness pressures of human development global warming.
from cholera and other bacterial have grown ever more powerful. Carbon and other greenhouse
infections fell dramatically, and gases are emitted from cars and
the river became much cleaner. Trees and climate change factories burning fossil fuels. Since
Human activity has always Forest has been especially hard-hit, 1958, American scientist Charles
altered the environment, but its mainly due to the dual demands Keeling’s measurements of
impact increased dramatically of lumber required for construction atmospheric CO2 have shown that
in the mid-18th century with the and fuel, and land cleared for CO2 emissions are increasing at
Industrial Revolution that began in agriculture and development. an ever-faster rate. While a minority
Britain, and spread to Europe, North An estimated 54,000 sq miles of scientists maintain that human
America, and beyond. The negative (140,000 sq km) of tropical rain activity is not responsible, climate
effects can be broadly divided forest—which contains the greatest change has warmed the continents.
into pollution, and destruction biodiversity—is cleared each year. The consequences, including trees
of resources and habitats. Scientists will never know how coming into leaf and flowers
THE HUMAN FACTOR 229
The International Charles Moore finds Camille Parmesan Maude Barlow wins
Dark Sky a mass of floating and Gary Yohe Canada’s highest
Association debris that becomes publish evidence for environmental
is founded in order known as the “Great the climate-change honor for her
to prevent light Pacific Ocean phenomenon of campaign for global
pollution. Garbage Patch.” “spring creep.” access to clean water.
1988 1997 2003 2008
1992 1999 2003 2014
Canada halts High mortality rates Research finds that Naomi Klein blames
its cod fishing in frogs are traced to ocean acidity large corporations
industry due viruses exacerbated has surged since for depleting natural
to excessively by the trade in live industrialization resources and the
depleted fish stocks. amphibians. began. climate crisis.
blooming earlier in spring, may caused by emissions of sulfur global population was 3.6 billion.
benefit some organisms but could dioxide and nitrogen oxide from By 2018, it had swollen to 7.6 billion,
prove disastrous for others. industrial chimneys. As a and although the growth rate
result, legislation to control the has slowed considerably, the
Toxic controls emissions was passed in the US ever-increasing consumption
The introduction of pesticides, such and Europe. After US chemists of natural resources has led to
as DDT, to increase crop harvests Frank Rowland and Mario Molina depleted stocks of wood, fossil
proved to be an environmental showed that chlorofluorocarbons fuels, minerals, and even fish. The
disaster: they eradicated useful (CFCs) destroy atmospheric ozone, collapse of the once bountiful cod
invertebrates as well as harmful the use of CFCs was banned fishery off Newfoundland in 1992
ones; they caused cancers in worldwide in 1989. highlighted the vulnerability of our
humans; and rendered birds of prey Light pollution, which affects food chain to overfishing and led
infertile. Rachel Carson’s 1962 book beach-nesting turtles, bats, and the Canadian government to
Silent Spring highlighted many of migrating birds, has proved harder impose an indefinite moratorium
these issues, and caused a partial to control. The International Dark- on fishing on the Grand Banks.
rethink of pesticide use. The work Sky Association is at the forefront Clean water is one of the most
of several other ecologists has of campaigns for environmentally fundamental requirements for
resulted in legislative controls to responsible lighting. society but almost 1 billion people
mitigate the environmental impact. do not have access to it. A lethal
When Gene Likens and his Diminished resources combination of climate change
team investigated why previously Garrett Hardin, an American and population growth in some
fish-rich lakes had died, they found ecologist, warned of the dangers developing regions threatens
that the culprit was acid rain, of overpopulation in 1968, when the to increase this number. ■
ENVIRONMENTAL
POLLUTION IS AN
INCURABLE
DISEASE
POLLUTION
232 POLLUTION
IN CONTEXT Effects of pollution on health

KEY FIGURE
Emma Johnston (1973–) The brain. Mental
development in
BEFORE children is delayed,
1272 King Edward I of and the brains of
adults are affected.
England bans the burning
of sea coal in London because
of the smoke it produces.
19th century Coal-burning The lungs. Air
pollution can
during Britain’s Industrial cause cancer,
Revolution stunts children’s and is linked
The heart. Pollution
to asthma.
growth and raises death rates increases the risk
from respiratory diseases. of heart disease
and stroke.
AFTER
1956 The Clean Air Act is The pancreas.
The reproductive
introduced in the UK, bringing Air pollution has
been linked to system. During
the thick smogs that plagued Type 2 diabetes pregnancy, the
fetus’s brain
its major cities to an end. in adults.
development can
be affected.
1963 The Clean Air Act
is passed in the US.
1972 The Clean Water Act
is fully ratified in the US.
1984 Toxic gas leaks from the
Union Carbide India factory
in Bhopal kill thousands and
injure many more. Polluted air and water cause the deaths of millions of
people every year. This illustration describes the specific
damage caused to different organs of the human body.
P
ollution comes in many be carried through air and water, thousands of years, indicates
forms, ranging from toxins affecting all life. Contaminants such that early humans generated air
in the air to trash at the as plastics can facilitate the invasion pollution from their fires. Analysis
bottom of the sea. Any substances of nonindigenous species, as of 2,500-year-old ice cores in
or forms of energy that spoil the discovered by Australian marine Greenland has shown evidence of
quality of the atmosphere, oceans, biologist Emma Johnston. There is air pollution from copper smelting
water, or soil are pollutants. They also a direct effect on human health: thousands of miles away, in the
may be chemicals or biological it is estimated that exposure to center of the Roman Empire.
contaminants (including human polluted air, water, and soil caused However, such impacts were
waste), products (such as plastics), 9 million premature deaths—one in on a small scale. With the start
or noise, light, or heat. The effects six of all deaths—in 2015. of the Industrial Revolution in
of pollution on life of all kinds Europe, air and water pollution
can be far-reaching, spreading Pollution through the ages became serious. Factory chimneys
thousands of miles beyond its Human-made pollution has a long pumped smoke out into the air;
original source. Pollution can history. The presence of soot on toxic chemicals poured into rivers.
spread through the food chain and the walls of caves, dating back Cities expanded quickly and had

See also: Pesticides 242–247 ■ Acid rain 248–249 ■ Light pollution 252–253 ■ A
plastic wasteland 284–285 ■ The water crisis 286–291 ■ Waste disposal 330–331
The “Great Stink”
By the early 19th century,
London’s Thames was the
centers fared little better: similarly
most polluted river in the
unsanitary conditions were recorded world. Industrial pollution and
in Berlin in 1870, for example. human effluent emptied into
In the United States, the first two it from thousands of drains.
cities to enact laws to ensure clean People complained, but the
Air pollution control air were Chicago and Cincinnati, in government did nothing. In
systems still lag behind 1881. By that time, the manure from 1855, the scientist Michael
economic development. 3 million horses pulling wagons in Faraday lambasted politicians
Bob O’Keefe North American cities was seeping for their inaction, to no avail.
into water supplies and producing However, they got the message
plagues of disease-causing flies. three years later, when a hot
As horses were gradually replaced summer contributed to the
by the internal combustion engine, “Great Stink” of 1858. The
smog from cars and trucks became Houses of Parliament, being
a major issue. London’s Great Smog adjacent to the Thames, were
badly affected, and legislation
no sanitation. The Thames River, of 1952, described as a “pea-souper”
was suddenly enacted in a
in London, was both the source for the color of the filthy air, killed mere 18 days.
of water for domestic use and the more than 4,000 people. Civil engineer Joseph
outlet for untreated human sewage. Bazalgette was commissioned
Disease spread, river fish were Air pollution to design a new sewage
wiped out, and the smell was The result of harmful substances system. It was based on six
sometimes unbearable. Other urban being released into the atmosphere, interceptor sewers, 100 miles
such as gases or small particles (160 km) long, which flowed to
called aerosols, air pollution can new treatment works. Most of
Of the world’s 20 worst cities for air London was connected to
pollution, 14 are in India. In Delhi, thick have natural sources, such as
volcanoes or wildfires, but is mainly it within a decade. Much of
smog in November 2017 reduced the
air quality to the equivalent of smoking caused by human activity. The main the sewage system is still in
50 cigarettes a day. air pollutants are emissions ❯❯ operation today, more than
150 years later.
This cartoon, published in Punch

magazine in July 1858, was entitled
“The Silent Highwayman.” People
at the time attributed the spread
of cholera to the bad river smells.
234 POLLUTION
The World Health Organization Pollutants entering
(WHO) estimates that nine out of the oceans
ten people worldwide are breathing
polluted air, causing widespread
illness and allergies. Furthermore, 10%
Pollution is one of the some aerosols, depending on the 20%
biggest problems we are composition and color of the 10%
facing globally, with horrible particles, block the amount of solar
future costs to society. radiation reaching Earth’s surface, 5%
Maria Neira thus having a cooling effect on the
5%
planet. Efforts to reduce air 20%
pollution can therefore make the
effects of global warming worse.
30%
Rivers, lakes, and seas
Surface water, groundwater, and the
from fossil-fuel-burning power oceans become contaminated by Air pollutants Offshore oil
stations, factories, motor vehicles, toxic chemicals from industry, from
Farmland runoff Industrial
the burning of wood and dung for chemical runoff from farmland, from
wastewater
heat and cooking fuel, and methane general trash such as plastics, and Sewage
from cattle, landfill sites, and from human waste. Maritime
Litter transportation
fertilized fields. Poor air quality Some rivers and lakes are
damages human health and crops, so polluted that they can support
and some fossil-fuel emissions cause no life at all, deprive communities In the oceans, the most acutely
acid rain, which has killed forests of freshwater and food, and carry destructive pollution has resulted
and fish in thousands of lakes. a risk of waterborne diseases, such from disasters involving oil tankers
as polio, cholera, dysentery, and and oil terminals. When the Exxon
typhoid. The WHO estimates that Valdez supertanker broke up on
Orcas may become extinct as a result
of PCB (polychlorinated biphenyl) 2 billion people worldwide are rocks off the coast of Alaska in 1989,
pollutants. The compound becomes drinking water contaminated with 11 million gallons (50 million liters)
more concentrated higher in the food human waste, resulting in the of crude oil were released into the
chain, and orcas are apex predators. deaths of 500,000 people a year. North Pacific. The oil smothered or
Emma Johnston Born in 1973, Australian marine She has also studied marine
biologist Emma Johnston was communities in the Antarctic,
interested in the oceans from an developed new biomonitoring
early age. She gained her Ph.D. in techniques, and advised
marine biology in 2002 and, in agencies on the management
2017, became the Dean of Science of estuarine biodiversity.
at the University of New South
Wales (UNSW), and Head of the Key works
UNSW’s Applied Marine and
Estuarine Ecology Lab, which 2009 “Contaminants reduce
investigates human impacts on the richness and evenness of
marine ecosystems. marine communities,”
Johnston discovered how Environmental Pollution
nonnative species can invade 2017 “Building ‘blue’: an eco-
waterways in coastal areas by engineering framework for
adhering to rafts of plastic foreshore developments,” Journal
pollution floating on the oceans. of Environmental Management
poisoned an estimated 250,000 in the 1920s. It can cause problems is a form of thermal pollution. It can
seabirds, 250 Bald Eagles, 2,800 sea for nocturnal wildlife, for example, kill fish and alter the composition of
otters, 300 harbor seals, and 22 because predator–prey relations are the food chain, reducing biodiversity.
killer whales. Billions of salmon and interrupted. Excessive noise can be Nuclear energy is sometimes
herring eggs also died. Further highly disturbing in cities, on flight viewed as “cleaner” than fossil-fuel
catastrophic damage followed in paths, and near factories and roads. energy, because it does not produce
1991, during the Iraq War, when But it also affects wildlife in subtler greenhouse gases, but it does result
Iraqi forces opened the valves of an ways. There is evidence that some in waste that remains radioactive for
offshore oil terminal and released at birds now sing at night because thousands or millions of years. The
least 380 million gallons (1,700 their song can be heard more industry also bears the inherent risk
million liters) into the Persian Gulf. clearly than during the day. of accidental damage. An explosion
The long-term effects of such Waste heat, too, can be at the Chernobyl nuclear power plant
disasters are still unfolding and damaging. When water from rivers in Ukraine in 1986 killed dozens of
have yet to be fully understood. or the sea is used as a coolant in people and spread radiation across
Much of our nondegradable factories or power stations, the hot Western Europe. The slowly
products ends up in the oceans. water that is returned to the source dwindling effects of contamination
Since the 1950s, around 8.3 billion on the ecosystem and human health
tons of plastic has been produced, are predicted to last a century.
of which only a fifth has been
recycled or incinerated. Each year, Mitigation measures
a staggering 8 million tons of Tackling the problem of pollution
plastic reaches the oceans, and is is a huge challenge, and involves
responsible for the deaths of huge In 2015, pollution both cleaning up existing pollution
numbers of marine animals. caused three times and making changes to reduce the
as many deaths as rate at which we add to it. Key
Intangible pollutants AIDS, tuberculosis, and aspects of this include replacing
Pollution in the form of energy, be malaria combined. fossil fuels with sustainable energy,
it light, noise, or heat, can be just Philip Landrigan more recycling and reuse, and the
as intrusive as physical waste or replacement of nondegradables
chemical emissions. Light pollution with degradable materials. This will
from buildings, streetlights, vehicles, take time and, ultimately, demands
and advertising billboards was first a fundamental shift in our culture
described as a problem in New York of consumption. ■
236
IN CONTEXT
GOD CANNOT
KEY FIGURE
John Muir (1838–1914)
BEFORE
SAVE THESE
1872 Yellowstone, in the states
of Wyoming, Montana, and
Idaho, is declared a national
park—the first in the world.
TREES FROM
AFTER
1948 The International Union
for Conservation of Nature
(IUCN), a partnership of
governments and civil society
FOOLS
organizations, is founded.
1961 The World Wide Fund for
Nature (WWF), initially known
as the World Wildlife Fund, is
formed, to protect endangered
ENDANGERED HABITATS species and habitats.
1971 The Man and the
Biosphere Programme (MAB)
is founded by the United
Nations, to promote sustainable
development. It has a global
network of Biosphere Reserves.
T
he origin of the movement
to conserve natural habitats
is usually credited to the
Scottish–American naturalist John
Muir, described as the “father of
the national parks.” He was one of
the first to realize that in order to
survive, wild places needed legal
protection. Of the many types of
natural habitat on Earth, some are
more fragile than others, but each
faces different threats, whether
anthropogenic (human-made) or
from natural causes, or both, and
many are critically endangered.
Habitats have, of course, always
been affected by destructive natural
events. Every year, lightning strikes
trigger large grassland and forest

See also: Human activity and biodiversity 92–95 ■ Biodiversity hotspots 96–97
■ Biomes 206–209 ■ Deforestation 254–259 ■ Environmental ethics 306–307
John Muir
Born in Scotland in 1838, John
Muir developed a passion for
nature as a boy. He moved
with his family to Wisconsin
at the age of 11. In 1867, he
had an accident in which he
lost his sight temporarily, after
which he “saw the world in a
new light.” An accomplished
botanist, geologist, and
Yosemite National Park was created glaciologist, Muir visited the
fires. Hurricanes and rivers in flood
in 1890, thanks to the efforts of John Yosemite Valley in California
can wreak havoc. Storm surges may
Muir. The park is famed for its glaciers, in 1868, and later determined
produce inundations of the sea, to preserve it from the scourge
waterfalls, and granite rock formations,
turning freshwater wetlands saline. such as the El Capitan monolith. of domestic sheep (which he
About 65 million years ago, the called “hoofed locusts”). In
Chicxulub meteor impact in Mexico 1903, Muir took President
produced a dust cloud so great that growth of cities, the development Theodore Roosevelt on a
it stopped sunlight from reaching of large-scale industry based on the guided tour through the
Earth’s surface. Plants struggled extraction of fossil fuels and raw Yosemite Valley, and their
to photosynthesize, and many materials, a growing agricultural three-day trip inspired
animals, including the dinosaurs, demand to feed more people, and Roosevelt to create the US
became extinct. conflict and war. All these have Forestry Service and, in
Nor is human influence an taken their toll on the natural world. 1916, to form the National
exclusively recent issue. Throughout Conservation Commission.
history, people have modified their Fragile ecoregions Until his death in 1914, Muir
continued to advocate for the
environment. Deforestation, for A concept that is now often used conservation of land such as
example, is not a new problem. In to identify the major habitat types Mount Rainier, which became
Europe, the clearance of forests for on Earth is that of the ecoregion— a National Park in 1899.
agriculture and construction began smaller than a biome, with a more
thousands of years ago, and a similar detailed gauge of biodiversity.
pattern followed in North America. Ecoregions are defined as large Key works
However, the impact of modern- units of land or water containing
1874 Studies in the Sierra
day humans on the environment is a geographically distinct mix of 1901 Our National Parks
unprecedented. In the past 200 species, natural communities, and 1911 My First Summer
years, the human population has environmental conditions. Some in the Sierra
exploded. This has fueled the rapid examples include deserts, tropical ❯❯
238 ENDANGERED HABITATS

absorb the greenhouse gas carbon
dioxide, thereby contributing to the
Natural factors can put Humans can put acceleration of climate change.
pressure on a habitat. pressure on a habitat.
Loss of coral reefs
Coral reefs are important ecoregions
and yet are especially endangered.
They support about 25 percent of
Pressures include the planet’s marine species, and
Pressures include are also nurseries for billions of fish.
deforestation, urban
storms, floods, wildfires, Two-thirds of the world’s reefs are
sprawl, mining,
volcanic activity, and
industrialization, under threat, and about a quarter
even meteor strikes.
pollution, and war. of them are likely damaged beyond
repair. Possibly the biggest threat
to coral reefs is increased acidity
caused by a greater uptake of CO2
from the atmosphere. This impedes
The habitat becomes endangered the ability of many sea creatures
to build their shells, and induces
coral “bleaching,” which is a step
on the way to the reef dying.
rain forests, temperate coniferous Once deforestation starts, the In addition, coral reefs are being
forests, lakes, mangrove swamps, problem quickly gets worse. When destroyed by overfishing, and by
and coral reefs. Of these, coral reefs rain falls on a forested slope, it is harmful practices such as cyanide
and tropical rain forests are under mostly absorbed by vegetation. and blast fishing, and bottom
particular threat from humankind. But when the slope is cleared, the trawling. Sediment resulting from
rainfall erodes the soil, making it coastal development blocks the
Rain forest clearance useless for agriculture and sunlight that reefs need. Chemical
Despite covering only 6 percent impossible to replant. It produces pollution, coral mining, and careless
of Earth’s land surface, tropical rain silt runoff into rivers and lakes, tourism all add to the burden
forests represent the greatest killing fish, and increases the risk on this highly sensitive habitat.
biomass of any terrestrial ecoregion, of flooding. The destruction of any
and are home to about 80 percent forest reduces its capacity to Wide-ranging impacts
of land species. Every year, some All over the world, diverse natural
54,000 sq miles (140,000 sq km) of habitats are critically threatened by
tropical rain forest is cleared—the human activity. Tropical deciduous
equivalent of a soccer field every dry forest is easier to clear than
second. Logging is carried out for rain forest, and on Madagascar,
firewood and construction materials, where dry forest was widespread,
and is also driven by the demand for less than 8 percent now remains. At
roads, settlements, and agriculture. one time, tallgrass prairie stretched
Globally, the rain forests that across the US Midwest, but only
are under greatest threat are in 3 percent of it is left: the rest has
West Africa, Central America, and been converted to farmland. Many
Southeast Asia. Indeed, only about wetlands have been drained for
30 percent of the lowland rain forest
in Borneo now survives . In the
Palm oil trees are being planted on
Amazon Basin, home to nearly a large scale in Indonesia and Malaysia,
one-third of the world’s rain forest, where this is one of the main drivers
much of the clearance is for of deforestation. Orangutans are among
agriculture, especially ranching. the species endangered as a result.

Protected areas
National parks, wilderness
areas, nature reserves, and
sites of special scientific
interest (SSSIs) are all types of
protected habitats. Within
these areas, interference with
the natural environment is
prohibited or limited by some
kind of legal framework. They
must cover a specified
expanse of land or sea, but
they vary greatly in size and in
the level of protection given.
Just over 10 percent of Earth’s
land is protected, but only 1.7
percent of the oceans; though
marine reserves are essential,
Wetlands and intertidal zones are
they require local and national
gases. Conservation is now
important for marine invertebrates and governments to agree on key
paramount, and ecologists work issues such as fishing rights.
migratory shorebirds, but in many parts to refine their understanding
of the world they have been drained for Marae Moana, the largest
industry and ports.
of the best ways to go about it. protected area on Earth, is
Appropriate measures depend 772,000 sq miles (2 million sq
on the situation, and range from the km) around the Cook Islands
agriculture or urban development; creation of protected reserves or in the Pacific Ocean. It is home
others are irreversibly damaged “corridors,” to link areas that have to sea turtles, at least 136
by pollution. Nutrient runoff from become fragmented, to projects species of corals, and 21
agricultural fertilizers has spoiled to recreate lost habitat. Sustainable whales and dolphins. The
many lakes and rivers. In many sources of fuel and timber for those biggest land reserve is the
countries, intertidal zones have who are otherwise dependant on Northeast Greenland National
been destroyed by the building forest wood are also important, as Park, which covers almost
of ports. Coastal development has is banning the trade in rain forest 386,000 sq miles (1 million sq
km) of ice sheet and tundra.
been largely responsible for the loss hardwood. Since the impact of
of 35 percent of mangrove swamps. habitat destruction is global,
In the tropics and subtropics, international agreements and
overgrazing by domestic animals cooperation are crucial. ■
such as goats has converted an
estimated 3.5 million sq miles
(9 million sq km) of seasonally dry
grassland and scrub into desert.
Halting the decline

The destruction of these habitats is In every walk with
not only a loss in terms of natural nature, one receives far
beauty and biodiversity, but also more than he needs.
creates serious problems for people: John Muir
for example, poorer water quality, Muskoxen are Arctic herd animals
declining fish stocks, crashes in whose numbers were severely
depleted in the 19th century by
populations of pollinators, flooding hunting. They now live on reserves
from increased rainwater runoff, in Alaska, Norway, and Siberia.
and faster buildup of greenhouse
240
WE ARE SEEING
THE BEGINNINGS
OF A RAPIDLY
CHANGING
THE KEELING CURVE
PLANET
T
he Keeling Curve, named more heat to be trapped, leading to
IN CONTEXT after Charles Keeling, an an overall increase in temperature
American scientist, charts and global climate change.
KEY FIGURE
the daily record of atmospheric
Charles Keeling (1928–2005)
carbon dioxide (CO2), measured in Measuring CO2 levels
BEFORE parts per million by volume (ppmv), Since the start of the Industrial
1896 The Swedish chemist in a series dating back to 1958. Revolution in the late 1700s, human
Svante Arrhenius is the first It shows two things: the natural activity has produced increasing
to estimate the extent to seasonal respiration of Earth and emissions of CO2. This is largely
which atmospheric CO2 could the year-on-year rise in atmospheric due to burning fossil fuels, while
increase Earth’s temperature. CO2. Atmospheric CO2 is significant forest clearance for agriculture and
because carbon dioxide is the most development has resulted in less
1938 Comparing historic important of the greenhouse gases, vegetation absorbing CO2 through
temperature data and CO2 which trap warmth in Earth’s photosynthesis. Many scientists
measurements, the British atmosphere. More molecules of CO2 once believed that excess CO2
engineer and scientist Guy and other greenhouse gases cause would be absorbed by the oceans.
Stewart Callendar concludes Others disagreed, but there was
that the increase in CO2 is little hard evidence either way.
responsible for the warming Charles Keeling was not the
of the atmosphere. first to propose a link between
atmospheric warming and CO2
AFTER emissions. Others had measured
2002 The European Space We were witnessing for CO2 levels but had produced only
Agency’s ENVISAT satellite the first time nature’s “snapshots” in time rather than a
begins to produce up to 5,000 withdrawing CO2 from the long-term dataset. Keeling knew
readings of greenhouse gases air for plant growth during that a long study was needed to
every day. summer and returning it prove the link. In 1956, he took up
each succeeding winter. a post at the Scripps Institution of
2014 NASA’s Orbiting Carbon Charles Keeling Oceanography in San Diego,
Observatory generates up to California, and obtained funds to
100,000 high-precision establish CO2 monitoring stations
measurements daily. at remote locations 9,843 ft (3,000 m)
up on Mauna Loa, Hawaii, and at
the South Pole. By 1960, Keeling

See also: Global warming 202–203 ■ The biosphere 204–205 Environmental
feedback loops 224–225 ■ Halting climate change 316–321
■
CO2 analysis in icecaps
Scientists can measure past
concentrations of carbon
dioxide by analyzing bubbles
of air trapped in Antarctic and
Greenland ice sheets. This
evidence indicates that there
have been several cycles of
variation over the past 400,000
years. These range from lower
readings in the most severe
glaciations—when glaciers
actually formed—to higher
readings during warmer,
interglacial periods. The
increase since the start of the
Industrial Revolution has been
matched by the average global
temperature. This has risen
by 0.13°F (0.07°C) per decade
since 1880 and 0.31°F (0.17°C)
Mauna Loa in Hawaii is an ideal site reflecting seasonal changes. During per decade since 1970.
for an atmospheric research station. The spring and summer in the Northern The Intergovernmental
high altitude and remote location of the Hemisphere, as new foliage takes
volcano ensure that the air is largely Panel on Climate Change
more CO2 out of the atmosphere, (IPCC) warns that unless the
unaffected by humans or vegetation.
the global concentration of the gas world's governments reduce
declines, reaching a low point in greenhouse gas emissions
was sure that he had a long-enough September. It increases again in dramatically, by the year 2100
series of records to detect a year- the northern autumn as leaves fall average temperatures could
on-year increase. and photosynthesis declines. Plant be around 7.7°F (4.3°C) higher
growth in the Southern Hemisphere than they were before the
Seasonal changes later in the year does not make up Industrial Revolution. Such an
Although South Pole funding ended for the loss, because most of Earth’s increase would cause both a
marked rise in sea levels and
in 1964, Mauna Loa has produced vegetative cover is in the north.
more extreme weather, which
data from 1958 onward. Plotted on a Ancient air bubbles trapped in would result in people having
graph, the measurements became polar ice cores reveal that, over the to abandon some regions of
known as the Keeling Curve. It is, past 11,000 years, average CO2 the world entirely.
in fact, a series of annual curves, concentrations were 275–285 ppmv,
but increased sharply from the mid-
19th century. In 1958, the level was
Mauna Loa CO2 record 316 ppmv. It rose steadily at a rate
(1958–2015) of 1.3–1.4 ppmv each year until the
CO2 concentration (ppmv)
400
mid-1970s, then increased by about
2 ppmv each year. By spring 2018 it
380 had hit 411 ppmv, almost 1.5 times
higher than preindustrial levels. ■
360
340 The Keeling Curve of steadily rising Bubbles in an ice core provide
CO2 levels is clearly shown on a graph a sample of the atmosphere going
320
plotting results from the continuous back centuries. Scientists measure
1960 1970 1980 1990 2000 2010
monitoring of atmospheric carbon the CO2 in the trapped air bubbles.
Years dioxide (CO2) at Mauna Loa, Hawaii.
THE CHEMICAL
BARRAGE HAS
BEEN HURLED AGAINST THE
FABRIC OF LIFE
THE LEGACY OF PESTICIDES
244 THE LEGACY OF PESTICIDES
A
rguably the most revered
IN CONTEXT and influential book ever
published on the subject
KEY FIGURE
of environmentalism, Silent Spring
Rachel Carson (1907–64)
garnered a huge amount of
BEFORE publicity when it was released in
1854 Henry David Thoreau’s 1962. It galvanized the fledgling
book Walden describes a social conservation movement, forced
experiment to live the simple legislative change, and, perhaps
life in tune with nature. It is most significantly, championed the
seen as an inspiration for the right of the public to question those
environmentalist movement. in power and hold them to account.
However, the author of this Spraying insecticide such as
1949 A Sand County Almanac ground-breaking work was far from DDT whether indoors or outside, has
by Aldo Leopold proposes a the typical “eco-warrior”—a term been—and in some places still is—a
that was unheard of when the book common method of controlling the
deep ecology of people living mosquitoes that transmit malaria.
in harmony with the land. was first published. On the
contrary, Rachel Carson was a
AFTER quiet, scholarly woman, with Boston Herald. It spoke about aerial
1970 The US establishes a masters degree in zoology and spraying of a mixture of fuel oil and
the Environmental Protection 20 years’ service as an aquatic a chemical compound named DDT
Agency (EPA). biologist in the United States. Most (dichloro-diphenyl-trichloroethane),
of all, she was an exceptional in the vicinity of her small bird
1989 Bill McKibben’s book
writer, able to fuse scientific fact sanctuary in Michigan. The
The End of Nature highlights
with compelling narrative. morning after the spraying,
the dangers of global warming. Huckins found several birds dead
2006 The documentary An Dying wildlife on her property and hoped that
Inconvenient Truth records Like many great and influential Carson might know someone in
former US vice president Al works, Silent Spring began in a very Washington who could stop further
Gore’s efforts to educate the personal way. In January 1958, spraying. Carson was outraged and
public about climate change. Carson’s friend Olga Huckins sent resolved to help. For more than a
her a letter that she had originally decade she had been aware of
tried to have published in the troubling incidents in which
Rachel Carson Born in 1907, Rachel Carson grew wrote books about marine
up on a farm in Pennsylvania, biology, most notably The Sea
where she developed a love of Around Us, which won the
nature. She won a scholarship to National Book Award, and was a
Pennsylvania College for Women national best seller. This success
and later gained a masters in enabled Carson to write full
zoology. Growing up in a land- time and she began work on
locked state, Carson dreamed of Silent Spring in 1958. In 1960,
the ocean; it became an enduring Carson was diagnosed with
passion, and she went to work as breast cancer; she died in 1964.
an aquatic biologist with the US
Fish and Wildlife Service (FWS). Key works
Carson wrote and published
many educational brochures and 1941 Under the Sea Wind
eventually became the US Fish 1951 The Sea Around Us
and Wildlife Service’s editor-in- 1955 The Edge of the Sea
chief. From 1941 onward, she 1962 Silent Spring

See also: Human activity and biodiversity 92–95 ■ Animal ecology 106–113 ■ The ecosystem 134–137 ■ A holistic view of
Earth 210–211 ■ Man’s devastation of Earth 299 ■ Environmental ethics 306–307
indiscriminate spraying of DDT had used to kill a wide range of insects,

been killing wildlife. Carson swiftly due to its pervasive action as a
approached the editor of the New nerve poison. It was used during DDT does not break
Yorker, E.B. White, suggesting that World War II to control insects down easily.
the magazine run a piece about the that destroyed vital food crops as
growing concern around synthetic well as those which transmitted
pesticides and their effect on malaria, typhus, and dengue fever
nontarget organisms. The editor to combat troops.
suggested that she write the article DDT proved cheap to produce,
herself. Reluctantly, Carson began highly effective, and at first
research on what she at first called appeared to pose no threat to DDT is fat soluble
“the poison book.” It went on to human beings. After the war, with and accumulates
shake the world. the chemical in plentiful supply, its in the body fat
use in agriculture was an obvious of animals.
The chemical future next step. With its wide range of
Silent Spring’s impact needs to be apparently safe applications, it
seen against the backdrop of the must have seemed like a panacea
time in which it was published. to farmers, who happily sprayed it
Although academics and scientists on their crops, often without the
had already voiced concerns about use of masks or protective clothing,
synthetic pesticides, the public because they did not fully appreciate
was oblivious to this issue. the powerful toxicity of this DDT is a broad-spectrum
Synthesized pesticides had dangerous chemical compound. poison, that affects not
been in use since the 1920s but had After DDT came a whole host of only the target pest
advanced significantly during World synthetic agrochemicals, including but other insects, fish,
War II, powered by military-funded aldrin, dieldrin, endrin, parathion, mammals, and birds.
research. During the 1950s, the malathion, captan, and 2,4-D. Used
popular notion was that they could in conjunction with fertilizers made
solve the world’s problems of out of surplus nitrogen that was no
famine and sickness by killing longer needed to make explosives,
pests that destroyed crops and these chemicals enabled the ❯❯
transmitted disease. Advertising
campaigns of chemical giants such
as Union Carbide, DuPont, Mobil, DDT can travel long
distances in the
and Shell spread this message to a
upper atmosphere.
huge audience. Silent Spring aimed
to challenge the received wisdom,
arguing that the so-called scientific No one since [Silent Spring]
progress enjoyed in post-war would be able to
America would come at a huge sell pollution as the
price for the environment. necessary underside
The most notorious of the of progress so easily.
pesticides, and the one most H. Patricia Hynes DDT causes
associated with Silent Spring, lasting harm
was DDT. It was first synthesized throughout the
in the late 19th century, but in food chain.
1939, Swiss chemist Paul Hermann
Müller realized that it could be
246 THE LEGACY OF PESTICIDES

DDT biomagnification in the food chain
A persistent poison
DDT (dichloro-diphenyl-
trichloroethane) belongs
to a group of pesticides
called organochlorides. It
kills insects on contact by
interfering with their nerve TERTIARY
impulses. The compound is fat 13.8 PPM
CONSUMERS
soluble and is deposited in the
tissues of animals exposed to
it, either directly or by eating
contaminated food. Repeated
exposure to DDT results in it
building up in the body fat and SECONDARY
2.07 PPM
becoming toxic. CONSUMERS
DDT also biomagnifies up
the food chain. Humans are
susceptible to poisoning from
regular exposure to DDT and PRIMARY
while the effects of small 0.23 PPM
CONSUMERS
amounts in the environment
are unknown, it has been
associated with cancer,
infertility, miscarriage, and
diabetes. It is now banned in PRODUCERS 0.04 PPM
western countries, but studies
carried out by the US Center
for Disease Control in 2003–4
Organisms higher in the food chain suffer from the impacts of DDT the most.
found DDT or its breakdown In producers, the poison only represents 0.04 ppm (parts per million), but the
product (DDE) in the blood of concentration increases with each step up the food chain. By the time tertiary
99 percent of people tested. consumers are involved, levels are high enough to have toxic effects.
intensification of farming. The stated that caution was essential

chemical age had dawned and by in the use of DDT because the true
1952, there were almost 10,000 impact of the product was not yet
separate new pesticide products fully understood. The following
registered with the US Department year, Fred Bishop, writing in the
A spray as indiscriminate of Agriculture (USDA). American Journal of Public Health,
as DDT can upset the stressed that DDT must not be
economy of nature… Raising awareness allowed to get into foods or be
Ninety percent of all Carson was not the first person to ingested by accident.
insects are good, and if notice the harmful effects of DDT. Various scientific studies and
they are killed, things go There were a few early dissenters, reports also raised concerns. For
out of kilter right away. including nature writer Edwin example, in 1945 the US government
Edwin Way Teale Way Teale, who warned that a published a study that found traces
spray with the indiscriminate of DDT in the milk of cows sprayed
impact of DDT could upset the with the chemical. It recommended
balance of nature. In 1945, the that farmers use “safe alternative
director of the US Fish and Wildlife substitute insecticides” to control
Service (FWS), Dr Clarence Cottam, flies and lice on cattle. Carson’s

Yorker—which had published a
serialization of the book. However,
Carson was prepared for this kind
of response. She knew the book
would be controversial and seen
They should as threatening by the chemical Man is part of nature
not be called industry. Therefore, as well as and his war against
insecticides meticulously tracking and recording nature is inevitably a
but biocides. her research—which had been war against himself.
Rachel Carson gained from government bodies, her Rachel Carson
contacts in research institutions,
and other reputable sources—she
also had the manuscript reviewed
by scientists and experts.
When suing Carson did not
work, the chemical companies
longstanding position as editor-in- launched a campaign to bring her protect the environment. The
chief at the FWS until 1952 meant into disrepute, stooping to personal Committee released a report
that she had access to a great many attacks such as depicting Carson entitled “The Uses of Pesticides,”
of these reports; she found them to as a “hysterical” cat-loving woman, which broadly supported Carson’s
be very disturbing reading. who was ill-equipped to write book. Inspired by Carson, activists
Since the research was rather such a book. The smear campaign continued to lobby government until
scattered and by no means backfired, merely increasing the in 1972, a decade after Silent Spring
accessible for the general reader, sales of Silent Spring. was first published, DDT was
Carson resolved to gather what banned in the US. Other countries
material she could find and present New policies followed, although some retain it to
it in a way that the ordinary non- Notable scientists supported control mosquitoes.
scientist could understand. As she Carson’s findings and US President The legacy of Silent Spring is
made progress with the writing John F. Kennedy invited her to greater than the banning of DDT.
of Silent Spring, it became clear testify before a Congressional It demonstrated to industry giants
to her that she had a moral duty to Committee in 1963. She called for and government the power of an
make the information public. As new policies that would serve to educated public. ■
well as documenting the hazards
of indiscriminate pesticide use,
Carson dared to suggest that the
chemical companies were putting
profits before people and that
the government might even be
colluding with them, knowingly
or otherwise, by failing to regulate
the industry effectively.
The response from the US
chemical industry was predictable.
At first, they tried to sue Carson,
her publishers, and The New
After DDT was banned in many

countries, Osprey populations—which
had declined significantly from the
1940s—began to recover. Ospreys had
eaten small animals affected by DDT.
248
A LONG JOURNEY
FROM DISCOVERY
TO POLITICAL
ACID RAIN
ACTION
T
he effects of acid rain on water quality and life forms in the
IN CONTEXT stone were noticed as long Hubbard Brook drainage basin in
ago as the 17th century New Hampshire. They discovered
KEY FIGURE
in England, and in Norway in the that the rainfall there was unusually
Gene Likens (1935–)
19th century. However, it was not acidic. Acidity, as expressed by pH
BEFORE until American ecologist Gene (potential of hydrogen), ranges from
1667 The corrosive effect of Likens carried out in-depth studies 0 (most acidic) through 7 (neutral),
polluted city air on limestone in an area of rural New Hampshire to 14 (least acidic). Most fish and
and marble is noted by the that the phenomenon came to be other aquatic animals fare best in
English diarist John Evelyn. properly understood. water with pH values of 6–8, but
From 1963 onward, freshwater Likens found values of 4—too acid
1852 British chemist Robert ecologist Likens and his team for fish, frogs, and the insects they
Angus Smith argues that studied the relationship between eat to survive. He set up monitoring
industrial pollution causes the stations around New England,
acidic rainfall that damages which showed that acid rain and
buildings. He is the first person snowfall were widespread in the
to call it “acid rain.” densely populated and heavily
industrialized northeastern states.
AFTER Likens’s systematic work persuaded
1980 The US Congress the US government to introduce
passes the Acid Deposition laws to control emissions of the
Act, undertaking an extensive chemicals responsible for acid rain.
18-year research program into
acid rain. Effects of acid rain
When fossil fuels are burned in
1990 An amendment to the
power stations and factories, sulfur
US Clean Air Act (passed dioxide (SO2) and nitrogen oxides
originally in 1963) establishes spew out of their chimneys.
a system that is designed
to effectively control
emissions of sulfur dioxide Acid rain had been wearing away
stonework—such as this statue in the
and nitrogen oxides.
churchyard of St. Peter and St. Paul,
Krakow, Poland—for hundreds of years
before the phenomenon was understood.

See also: Endangered habitats 236–239 ■ Pesticides 242–247 ■ Deforestation
254–259 ■ Depletion of natural resources 262–265 ■ Ocean acidification 281
Fossil fuels burned These gases react

by industry emit with water in the
sulfur dioxide and lower atmosphere to
nitrogen oxides produce sulfuric acid
into the air. and nitric acid.
Gene Likens
Plants and
animals The acids fall as rain Likens was born in Indiana
cannot live in into lakes and rivers, in 1935. After earning a Ph.D.
affecting the pH balance in zoology from University of
the acidic water of the water.
and die off. Wisconsin, he was appointed
assistant professor at
Dartmouth College. In 1963,
with fellow scientists F.
Herbert Bormann, Noye
Spreading through the lower By the early 1970s, thousands of Johnson, and Robert Pierce, he
atmosphere, these gases react with lakes in Scandinavia had lost their began research into the water,
minerals, and life forms in the
water to produce dilute sulfuric fish and were virtually dead. By 1984,
Hubbard Brook basin. In
acid (H2SO4) and nitric acid (HNO3). Brooktrout Lake and others in the
1968, his studies recorded
These weak acids fall as rain and Adirondack Mountains, New York, the widespread prevalence
enter rivers and lakes, making them were devoid of fish. Acid rain also of acid rain, the product of
more acidic. Increased acidity leaches harmful aluminum from the emissions from factories in the
stresses animals and plants. Water soil, and acidic clouds and fog harm Midwest. The team’s work in
snails disappear, fish eggs fail to plants, reducing their ability to the area over many years was
hatch, and insects and the frogs photosynthesize, leading to death. described as one of the world’s
that eat them die. Eventually, lakes most thorough studies of how
will not support any life. Emission control air pollution and land use has
In the 1970s and 1980s, other areas shaped a drainage basin.
badly affected by acid rain Likens’s work on deforestation,
included the “Black Triangle” of land use, and sustainability
Czechoslovakia, Germany, and led to a change in policy by
Poland, where large areas of forest the US Forestry Service. It also
helped shape the amended
died. Thanks to Likens’s work,
We experienced eight Clean Air Act in 1990. Likens
stricter controls were brought in was awarded the National
years of denial, but after 1990. Scrubber systems that Medal of Science in 2001.
that’s not unusual in extract SO2 were fitted to power
environmental issues. station chimneys with great Key works
Gene Likens success. Emissions of the gas were
cut by almost half in the US, and by 1985 An Ecosystem Approach
two thirds in Europe. Fish began to to Aquatic Ecology: Mirror
return to lakes and rivers. However, Lake and its Environment
the problem of acid rain still blights 1991 Limnological Analyses
parts of Russia, China, and India. ■
250
A FINITE WORLD
CAN SUPPORT ONLY
AOVERPOPULATION
FINITE POPULATION
I
n 1968, two scientists in the extinction. Hardin himself proposed
IN CONTEXT US issued dire warnings about a controversial solution to the
overpopulation. Ecologist problem of overpopulation, arguing
KEY FIGURE
Garrett Hardin predicted that that the government should deny
Garrett Hardin (1915–2003)
Earth’s resources would soon be welfare assistance to people who
BEFORE used up and environmental damage bred “excessively,” to prevent
1798 Thomas Malthus would increase. In The Tragedy of further births. Biologist Paul Ehrlich
forecasts that continued the Commons, he cited examples similarly advocated population
population growth will exhaust of several major global crises that control in The Population Bomb,
global food supplies by the had been caused by overpopulation: with warnings that human numbers
mid-19th century. the destruction of fish stocks by would soon reach a point where
overfishing; the draining of lakes mass starvation would ensue.
1833 In Two Lectures on the by over-extraction of groundwater
Checks to Population, British for irrigation; deforestation; pollution Growth and decline
economist William Forster of air, land, and sea; and species For most of human history, the
Lloyd discusses overpopulation, world’s population had grown only
using the example of common slowly. It began to increase more
land, which is less productive rapidly in Western Europe and the
if too many cattle graze it. United States in the early years
of the Industrial Revolution, when
AFTER British economist Thomas Malthus
1974 A United Nations warned of a future famine. His
conference in Bucharest fears, however, proved premature
creates the UN’s first World because food production increased
Population Plan of Action. more quickly than many expected.
Life expectancy also fell in the new
2013 British social geographer industrial cities, due to infectious
Danny Dorling outlines in diseases. It rose again with better
Population 10 Billion why it is
unlikely the world’s population
will ever reach that number, Ragpickers Court (1879) by William
Allen Rogers shows a poor Italian
contrary to UN estimates. neighborhood in New York City. Such
overcrowding allowed diseases to
spread through poverty-stricken areas.

See also: Human activity and biodiversity 92–95 The Verhulst equation
164–165 ■ Depletion of natural resources 262–265
■
■ Urban sprawl 282–283

China’s one-child
family planning policy
Until the 1960s, China
World population growth, 1750–2100 encouraged families to have
2.1% 11.2 Billion as many children as possible,
2% and the population rose
9.8 Billion
from 540 million in 1949 to
1.8%
940 million in 1976. However,
1.6% 8.6 Billion the government soon became
1.4% concerned about the demand
7.6 Billion
on resources. In 1978, scientist
1.2%
and politician Song Jian
1% calculated that China’s ideal
4.5 Billion
0.8% population was between 650
3 Billion and 700 million people, and in
0.6%
2 Billion
1979, his projections led the
0.4% 1.65 Billion government to create a new
0.2%
0.9 Billion policy limiting couples to one
0.1% child per family.
0%
This one-child policy was
1760 1780 1800 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 2040 2060 2080 2100
enforced more strictly in urban
This graph plots a comparison areas than in the countryside;
between the annual growth rate Annual growth rate as a percentage in some regions a second child
of the world population and the total of the world population was permitted if the first was
population in absolute numbers. a girl. In the cities, however,
The data for the years after 2017 World population in numbers women were forced to abort
is a projection. second children, and in 1983
alone, 21 million women were
forced to undergo sterilization.
healthcare and nutrition, cleaner still growing quickly in some parts The policy was relaxed in 2015,
water, and more rights for workers. of the developing world, but the but the government still only
By 1924, there were 2 billion people trend is not as rapid as it once was. allows two children per family.
in the world, and by 1960 there It took just 11 years for the world’s
were 3 billion, with most growth population to rise from 6 billion
occurring in the developing to 7 billion, but the increase to
countries of Latin America, Africa, 8 billion is forecast to be 13 years,
and South and East Asia. then another 25 years to reach
9 billion. The UN forecasts a peak
A slowing birthrate of 11.2 billion in 2100.
In Europe and North America in Despite the slowing growth,
the 20th century, wider access to challenges remain. In 2009, a UN
birth control, better education, and report warned that the world would
more women entering the labor need to produce 70 percent more
market resulted in lower birth rates. food by 2050 to feed its extra
This phenomenon is now being population, thereby putting more
replicated for women everywhere. pressure on land, water, and energy
Although the world’s population resources. Future population
passed 4 billion in 1974, 5 billion in growth is also likely to aggravate A 1994 poster of a smiling mother
1987, 6 billion in 2000, and 7 billion many environmental problems, and daughter promotes China’s
one-child policy. Many baby girls
in 2011, the annual rate of increase such as pollution, and rising levels were abandoned or killed so that
peaked near the end of the 1960s at of atmospheric greenhouse gases, their parents could try for a son.
2.5 percent a year. Populations are fueling global climate change. ■
252
DARK SKIES
ARE NOW
BLOTTED OUT
LIGHT POLLUTION
IN CONTEXT
KEY FIGURE
Franz Hölker
BEFORE
1000 ce The first organized
system of street lighting
(by oil lamps) is introduced
in Muslim Spain.
1792 Scottish-born engineer
William Murdock invents the
gas light. Over the next half
century, many cities introduce
gas street lighting.
1879 American inventor
Thomas Edison demonstrates
the first commercially viable
A
electric light bulb. ccording to some ecologists, A map of light pollution across
light pollution—the amount North America (white and red indicate
1976 High-brightness, where it is highest, black where it is
of artificially generated light
high-efficiency, LED lights lowest) explains why 99 percent of
in the world—could be the most
are introduced. Americans cannot see the Milky Way.
damaging pollutant of all. Around
AFTER 80 percent of humanity lives under
2050 The date by which skies saturated with light. In 2017, intense in industrializing countries
Hölker and others predict that, a major German study of light in South America, Africa, and Asia,
with the global population set pollution, carried out by ecologist but it also continues to increase
to exceed 9 billion, Earth’s Franz Hölker and others using in the already well-lit countries
total illuminated area will have satellite data, showed that the area of Europe and in the US.
doubled since 2016. of Earth illuminated artificially Astronomers were among the
grew 9 percent between 2012 and first to notice light pollution because
2016. The brightening is most it interfered with their ability to see

See also: Environmental feedback loops 224–225 Spring creep 274–279
■ Man and the Biosphere Programme 310–311
■
The effect on turtles
Light pollution is a major
problem for nesting sea
Artificial light can also damage
turtles, which lay their eggs
birds’ immune systems. Studies on land because the embryos
have found that house sparrows breathe through the
infected with the West Nile virus permeable shells. Females
carried the virus twice as long when need dark, sandy beaches for
Dark areas are being lost kept under dim light than when kept their nests, and will go
in places where nocturnal in the dark—doubling the time in elsewhere if there are bright
animals, insects, and plants which mosquitoes could bite them lights from beach resorts,
have adapted to darkness and pass on the virus. street lights, or housing. If a
over billions of years. Ill-effects on animals can have whole stretch of coastline is
Franz Hölker a knock-on effect on plants. When illuminated, they may lay their
moths, which are attracted to light, eggs in inferior habitats or
are repeatedly drawn to artificial even deposit their eggs at sea,
sources, not only can they be killed where their offspring will die.
Such problems may be the
by exhaustion (because the light is
reason for the reduction in sea
never extinguished), or by the heat
turtle populations. Scientists
generated, but they also become believe that hatchlings move
celestial objects in the night sky. more vulnerable to predators, which toward the brightest light. In
In 1988, American astronomers Tim can spot them more easily. natural conditions, this will be
Hunter and David Crawford founded The decline in moth numbers moonlight shining on the
the International Dark-Sky has a knock-on effect on the plants ocean, but if there is artificial
Association to protect the night that they help pollinate, which then lighting inland, the hatchlings
skies from light pollution. It was affects seed yield. In some places, wander toward that and get
the first organization of its kind. seed yield has declined by as much run over by traffic, eaten by
Since then, studies have as 30 percent. Researchers who predators, or caught in fencing.
examined the effects of light studied a Swiss flower meadow Solutions include getting
pollution on plants and animals, under street lights found that people and businesses to turn
which rely on the cycles of light nocturnal visits from pollinators off lights at night or use
and dark to govern life-sustaining declined by two-thirds. ■ “turtle-safe” lighting, which is
virtually invisible to turtles.
behaviors such as nourishment,
sleep, protection from predators,
and even reproduction. Such
research reveals a raft of ill-effects.
One study showed that trees in
Europe are budding more than a
week earlier than they were in the The solution is simple—
1990s; this alters their period of turn off unnecessary lights,
growth, and may mean that they fail use only the amount of light
to drop their leaves and fruit and needed for the task at hand,
enter the dormant phase in time to and shield all lighting
avoid damage over the winter. so it shines down
where it is needed.
Vicious circle Tim Hunter
Light pollution also has a detrimental
effect on animals. Lights on tall Olive ridley sea turtle
hatchlings make their way toward
towers, for example, draw migrating the sea at Boca del Cielo Turtle
birds, causing them to crash into Research Station, Mexico.
the towers and into power lines.
I AM
FIGHTING FOR
HUMANITY
DEFORESTATION
256 DEFORESTATION
D
eforestation is the removal
IN CONTEXT of forest or woodland for
conversion to nonforest
KEY FIGURE
Chico Mendes fought use. This can be conversion to
Chico Mendes (1944–88)
to save the tropical agricultural land, including cattle
BEFORE rain forest in Brazil. ranches, or development for housing,
1100–1500 Temperate forest industry, or transportation. Forest
is cleared across large parts may be degraded without being
of western and central Europe. destroyed completely, when valuable
mature trees, such as teak, are
1600–1900 Forests are cut selectively logged or some trees are
down in North America to cut down to create a road. This can
make room for agriculture. have a disproportionate negative
effect on the biodiversity of the
Late 1970s Tropical rain forest forest, even though most trees are
clearance, mostly for ranching, His local actions left standing. Another form of
accelerates dramatically. helped reduce global deforestation is the clearance of
AFTER CO2 emissions. primary forest and its replacement
2008 The UN launches its with monoculture plantations, such
Reducing Emissions from as palm oil, as has happened
Deforestation and Degradation extensively in Indonesia.
(REDD) incentive program. Deforestation can impact all
kinds of forest habitat, but tropical
2010 The US converts $21 m rain forest—tropical moist broadleaf
(£16 m) of Brazil’s debt into a forest that grows between the
fund that will protect Brazil’s Mendes Tropic of Cancer and the Tropic of
coastal rain forest. realized that he Capricorn—is the most severely
2015 The UN Paris Agreement had had a global

sets targets for planting trees effect: “I am fighting Polluting smoke swirls up as rain
to offset the threat of climate for humanity.” forest burns to make way for agriculture
in Brazil. It is estimated that Brazil
change and global warming. clears 2.7 million acres (1.1 million
hectares) of rain forest a year.
By felling trees … men

bring upon future
generations two calamities
at once: want of fuel and
scarcity of water.
Alexander von Humboldt
19th-century German explorer

See also: Biodiversity and ecosystem function 156–157 ■ Climate and vegetation
168–169 ■ Global warming 202–203 ■ A holistic view of Earth 210–211
affected. Concern for the rain forest wood for shipbuilding from the
was first raised in the 1970s when Baltic nations and New England
activist Chico Mendes—who went in the US.
on to become a founding member of Primeval forest clearance
Brazil’s National Council of Rubber accelerated globally between 1850
Tappers—called on the Brazilian and 1920, with the biggest losses in
government to establish forest North America, the Russian empire,
reserves, from which local people and South Asia. In the 20th century,
could extract natural products, the focus shifted to the tropics,
such as nuts, fruits, and fibers, especially to tropical rain forest,
sustainably. Mendes’s campaign, half of which has been destroyed Chico Mendes
which eventually cost him his life, since 1947, with the proportion of
highlighted the ecological damage the land that it covers having fallen Born in 1944, the son of one
wreaked by forest clearance. from 14 percent to 6 percent. of the 50,000-strong “Rubber
It is estimated that an area Army” who tapped rubber for
Human need equivalent to 27 soccer fields use in the Allied war effort in
The human race has used trees from is lost from forests globally each World War II, Mendes started
its earliest days. In Neolithic times, minute. Some regions have been work as a rubber tapper at the
they were cut down for fuel and hit harder than others. In the age of nine. Influenced by
to construct shelters and fencing. Philippines, for instance, 93 percent priests from the progressive
Five-thousand-year-old stone axes of tropical broadleaf forest Liberation Theology
for chopping wood have been has been removed; 92 percent movement, he helped found a
branch of the Workers’ Party
found, as well as ax factories from of Atlantic forest in Brazil has gone;
and became leader of the
the same era in Europe and North 92 percent of temperate coniferous
Rubber Tappers’ Union.
America. During the Middle Ages, forest in southwest China has As large areas of Brazil’s
however, as human populations disappeared; and 90 percent of dry rain forest were cleared to
expanded rapidly in western Europe broadleaf forest in California has make way for cattle ranches,
between 1100 and 1500, extensive been cleared. Mendes publicized the
deforestation took place. Forests tappers’ fight to save the
were cleared to make way for Effects on biodiversity forest. He went to
agriculture, and wood was used to Recent estimates suggest that Washington, D.C., to persuade
build homes and boats, and to make almost half of all forest clearance Congress and the World Bank
bows, tools, and other implements. is carried out by subsistence ❯❯ that cattle-ranching projects
Trees were cut down on an should not be funded. Instead,
industrial scale in central Europe hr proposed that forest areas
and England to produce charcoal, be protected as “extractive
which became an important fuel reserves”—public land
managed by local communities
(until replaced by coal) because
with the right to harvest forest
it burns at higher temperatures products sustainably. Cattle
than wood. An early example We are unable to
ranchers saw his movement as
of sustainable production was remain silent in the face a threat, and one, Darcy Alves,
practiced in England, where many of so much injustice. shot him dead in 1988. After
woods were managed as coppices Chico Mendes his death, the first of many
whose trees were partially cut such reserves was established,
back and then allowed to regrow covering 2.5 million acres
to create a cyclical supply of (1 million hectares) of forest
charcoal. Even so, by the 17th around Xapuri.
century England had to import
258 DEFORESTATION
farmers, and a third by commercial humans—not least because most
interests. Urban development, new medicines are derived from
logging for the best-quality lumber, plants, and so the eradication of
mining and quarrying, and trees the rain forest’s rich store destroys
cut for firewood account for any potential cures for disease.
remaining deforestation. In every I became an ecologist Rain forests, together with all
case, the environment suffers. long before I had ever other trees and woodland, also
Biodiversity is particularly impacted, heard the word. act like a sponge for rainfall.
because only a small number of Chico Mendes Tree roots drink up moisture and
mammal, bird, and invertebrate limit surface runoff. When forest
species can live on grassland or a is cut or burned, the soil is leached
palm oil plantation, and even fewer of many of its nutrients. If it covers
in industrial or urban settings. a slope, the soil will wash away,
Human conflicts also blight forest, leaving the land unfit for growing
the worst example being the Agent any kind of plants. Deep gullies
Orange chemical used to defoliate two-thirds of the world’s plants and may undermine trees that have
trees during the Vietnam War. animals live in this environment. not been cut, bringing them down.
Between 1.5 million and 1.8 million After heavy rains, catastrophic
The rain forest species—mostly insects, followed mudslides, which happen with
Destruction of the rain forest by plants and vertebrates—have increasing frequency, sweep down
poses a severe threat to global already been identified in rain the slope, destroying everything
biodiversity because it has been forests, and many others have yet in their path—including human
estimated that between half and to be discovered and described. In settlements. In May 2014, for
Borneo, Indonesia, for example, an example, heavy rainfall on the
area of just 0.2 sq mile (0.5 sq km) deforested slopes of the Caribbean
Replacing trees with human
settlements destabilizes the soil on may contain more species of tree island of Hispaniola caused
slopes, and mudslides, such as this than the combined landmass of mudslides and floods that killed
catastrophic event in Sierra Leone Europe and North America. Such more than 2,000 people. Conversely,
in 2017, are more likely to occur. biodiversity is vitally important to in extended periods of dry weather,

Deforestation in the Brazilian Amazon Reforesting
the Amazon
30000 About 17 percent of rain forest
in the Amazon Basin has
AREA DEFORESTED, IN SQ KM
25000 been lost since the mid-1970s.

At the United Nations Paris
20000 Climate Summit in 2015,
Brazil pledged to restore
15000 nearly 30 million acres (12
million hectares) by the year
10000 2030. In 2017, Conservation
International, in partnership
5000
with the Brazilian government,
launched the area’s biggest
0 reforestation program to date,
1990 1995 2000 2005 2010 2015 under which Amazonas state
will gain 73 million trees
The depleting rainforest cover in the Amazon Basin is a through seeding and planting.
global concern. The land is now being stripped of trees at a Local communities are
rate of 8,000 sq km (3,000 sq miles) annually. being enrolled to implement
the program, using a
technique called muvuca. This
exposed soil dries out faster than for conserving forest, and the involves spreading the seeds
tree-covered areas, making it more creation of extractive reserves, of more than 200 native forest
prone to wind erosion. where local people can harvest species over every square yard
products sustainably. of land. Less labor-intensive
Fueling global warming Globally, alternative sources than traditional tree-planting,
Burning wood or forests adds of fuel need to be found, along the method can reforest land
carbon dioxide (CO2) to the with new ways to develop less quickly, delivering around
atmosphere. By contrast, living land-hungry forms of agriculture. 6,000 plants per acre. In
plants of all kinds reduce CO2, A few nations are taking the lead addition to the seeding
as they absorb carbon, taking up in reforestation programs. For program some planting will
the greenhouse gas to perform example, a project in which people enrich secondary forest and
photosynthesis, thus countering from 500 villages have planted 150 return pasture land to forest.
the damaging impact of human million mangrove trees on the coast
activities. Globally, forests suck up of Senegal will restore mangrove
2.65 billion tons (2.4 billion tonnes) forests to boost fishing and shield
of CO2 every year. Environmentalists rice paddies from the influx of salt
and climatologists worry that water. The Chinese aimed to plant
removing large tracts of tropical 16.3 million acres (6.6 million
forest could be disastrous. hectares) in 2018, equal to the area
of Ireland; in 2000, the proportion of
Reforesting Earth China covered by forest had fallen
Currently, about 31 percent to 19 percent, but the target is to
of Earth’s land surface is covered increase this to 23 percent by 2020
by forests, but that figure is rapidly and 26 percent in 2035. ■
decreasing in some parts of the
world. However, there are regions,
The first African woman to win
including Europe, where forest a Nobel Peace Prize (2004), Wangari
areas are gradually expanding. Maathai initiated a community-based
Measures to restrict deforestation tree planting program to reverse
include payments to communities erosion and desertification in Kenya.
260
THE HOLE IN THE

OZONE LAYER IS A
KIND OF
OZONE DEPLETION
SKYWRITING
I
n 1982, a team of scientists in spring. Over Antarctica, ozone
IN CONTEXT working for the British Antarctic measurements have been down
Survey (BAS) discovered that by 70 percent compared with 1975.
KEY FIGURE
ozone levels above the Antarctic had Over the Arctic, levels have fallen
Joseph Farman (1930–2013)
fallen dramatically. Ozone (O3, a by nearly 30 percent. This effect
BEFORE colorless gas in the stratosphere, became known as “the ozone hole,”
1974 American chemists 12–18 miles (20–30 km) above Earth’s although it is better described as
Frank “Sherry” Rowland and surface, forms the “ozone layer,” a “the ozone depression,” since it is
Mario Molina suggest that protective shield that absorbs most a thinning of the ozone layer rather
chlorofluorocarbons (CFCs) of the Sun’s ultraviolet (UV) radiation. than a complete hole.
destroy atmospheric ozone. Without it, more of the Sun's harmful
radiation would reach the surface. Antarctic discovery
1976 The US National Since the mid-1970s, there has British geophysicist Joe Farman
Academy of Sciences declares been a 4-percent decrease in the was one of the team who made
that ozone depletion is a reality. amount of ozone in the stratosphere. the discovery in 1982. BAS teams
An even bigger decrease has been had been collecting atmospheric
AFTER seen above the poles, particularly data at the Halley Research Station
1987 The Montreal Protocol in Antarctica since 1957. Their
on Substances that Deplete work was poorly funded, and they
the Ozone Layer, a global relied on dated instruments such as
treaty to phase out CFCs and the Dobson meter—a rudimentary
similar chemicals, is agreed. machine that worked properly only
Joe Farman [made] when wrapped in a duvet.
1989 Montreal’s worldwide
one of the most important When Farman first noticed the
ban on the production of CFCs
drop in ozone levels, he found it
comes into effect (ratified by geophysical discoveries of
hard to believe, and thought there
the EU and 196 states to date). the 20th century. must be a problem with his Dobson
John Pyle and Neil Harris meter. He ordered a new instrument
2050 The year in which ozone Atmospheric scientists,
over the Antarctic is predicted University of Cambridge for the next year—and it recorded
to return to pre-1980 levels; an even bigger dip. The following
however, other harmful year, the dip was bigger again.
emissions may delay recovery. The year after, his team took their
measurements 620 miles (1,000 km)
from Halley. Again, there was a

See also: Global warming 202–203 ■ Environmental feedback loops 224–225 CFCs
■ Pollution 230–235 ■ The Keeling Curve 240–241 ■ Environmental ethics 306–307
Chlorofluorocarbons (CFCs)
are chemicals made up of
carbon, chlorine, and fluorine
atoms. They are non-toxic,
nonflammable, and extremely
stable. Their low reactivity
makes them very useful, but
is also the reason why they
are so destructive. They can
survive for over 100 years,
which gives them time to
diffuse into the stratosphere.
There, they are broken down
by the intense UV light to
release chlorine, which reacts
with ozone to form oxygen.
CFCs were first produced
in 1928, and were used as
coolants for refrigerators. They
were later used in a wide
range of aerosol products, for
example insect sprays, hair
conditioners, and spray paints.
The replacements for CFCs
were hydrochlorofluorocarbons
(HCFCs), which also deplete
the ozone layer, although to
a much lesser extent, and
hydrofluorocarbons (HFCs).
HCFCs will be phased out by
A NASA image of the “ozone hole” scientists Frank Rowland and 2020. HFCs do not harm the
over Antarctica in 2014. The blue area ozone layer at all—but they
Mario Molina. They had concluded
shows where there is least ozone. The are very powerful greenhouse
that gases containing chlorine—
amount of ozone in Earth’s stratosphere gases, and so in 2016 it was
overall has stabilized since about 2000.
including the chlorofluorocarbons agreed that, from 2019, they
(CFCs) used in aerosol sprays and too would be phased out.
halogen refrigerants—were, in
large dip. Farman decided it was the presence of UV light, reacting
time to publish, and a paper written with ozone in the stratosphere and
by him and his colleagues Brian breaking down the gas. A few
Gardiner and Jon Shanklin appeared countries, including the US, banned
in the journal Nature in 1985. the use of these products, but most
were yet to be convinced.
Reaction and response When ozone levels continued
Most scientists greeted Farman’s to fall throughout the 1980s, opinion
discovery with alarm: the potential gradually changed. Consequently,
increase in UV radiation would make in 1987, the Montreal Protocol for
skin cancers, cataracts, and sunburn a global ban was agreed. The ozone Aerosol products such as insect
far more prevalent. layer is showing signs of recovery, repellents were widely available
from the 1950s. The damaging
What could be done? One and it is hoped that by 2075, effects of the CFCs they contained
reason for ozone depletion had been stratospheric ozone will return were not known until the 1970s.
identified in 1974 by American to 1975 levels. ■
262
IN CONTEXT
WE NEEDED
KEY FIGURE
Naomi Klein (1970–)
BEFORE
A MANDATE
1972 The UN’s Conference on
the Human Environment calls
for an international approach
to environmental protection.
FOR CHANGE
1980 The World Conservation
Strategy, launched in 35
countries, introduces the
concept of sustainability.
1992 At the UN Earth Summit
DEPLETION OF NATURAL RESOURCES in Rio de Janeiro, member
states produce “Agenda 21,”
which outlines plans for
managing resources in the
21st century.
AFTER
2015 The UN Sustainable
Development Summit sets out
17 sustainable development
goals and launches a bold
global agenda, adopted by
193 member states.
I
n This Changes Everything
(2014), Naomi Klein railed
against the way that
governments and corporations
deplete natural resources. “Ethical
oil,” she maintains, is not just a
contradiction in terms, “it’s an
outrage.” A Canadian citizen,
Klein has campaigned against the
exploitation of the Athabasca tar
sands, the largest of three major
oil sand deposits in western
Canada. The oil sand deposits lie
under thousands of square miles
of coniferous forests. The open-pit
extraction of oil from tar sands
is particularly harmful to the
environment. Vast acres of forest
are cleared, and ponds of pollutants

See also: Deforestation 254–259 ■ Overfishing 266–269 ■ The water crisis 288–291
■Humankind’s dominance over nature 296 ■ Human devastation of Earth 299
Naomi Klein
Born in Montreal, Canada, in
1970 to politically active
parents, Klein developed a
sophisticated understanding
of the way the world works
while still young. Her first job
was on a Toronto newspaper,
The Globe and Mail. Her debut
book No Logo, criticizing
Extracting crude oil from Canada’s in a way that was sustainable and globalization and corporate
tar sands is notoriously harmful to fair for all nations around the world. greed, was a bestseller. Her
the environment. It accounts for a second, The Shock Doctrine,
Key areas in the struggle for a
tenth of Canada’s annual greenhouse attacked neoliberalism. Klein
sustainably managed Earth are
gas emissions. then began campaigning
use of fossil fuels, deforestation, against corporate interests
and water management. taking priority over the
are left behind. These can leak into Five years later, at the 1992 environment and the interests
the land, rivers, and groundwater, Earth Summit in Rio de Janeiro, of humanity. Her book This
killing fish, migrating birds, and 172 nations signed environmental Changes Everything was later
other animals. resolutions. Among them was made into a film. Klein’s many
Agenda 21, a plan for governments campaigns included a protest
Global action to work together to protect natural against the construction of the
By the 1980s, the environmental resources and the environment. Keystone XL pipeline—a
effects of industrialization and However, implementing changes symbol in the battle against
depletion of the Earth’s resources has proved difficult, and subsequent fossil fuel use and climate
were already becoming a matter of Earth summits have called for change. In November 2016,
concern. The United Nations (UN) better international cooperation she was awarded Australia’s
Sydney Peace Prize.
created a World Commission on in order to achieve goals set.
Environment and Development,
Key works
which published a report in 1987 Peak oil
called “Our Common Future.” Fossil fuels are among the world’s 2000 No Logo
Contributing experts, including most highly prized resources. 2007 The Shock Doctrine: The
scientists, agriculturalists, foreign People have become increasingly Rise of Disaster Capitalism
ministers, technologists, and reliant on oil, squandering it to 2014 This Changes
economists, made it clear that create a lifestyle that is ultimately Everything: Capitalism vs
the future of humans relied on unsustainable. The oil crises of the The Climate
balancing ecology and economics 1970s highlighted how dependent ❯❯
264 DEPLETION OF NATURAL RESOURCES

yet to be discovered. In the early loss of forests worldwide directly
21st century, new dates were given, causes 15 percent of global
some extending the timeline for greenhouse gas emissions.
oil to 2030 or beyond. In 2011, Rain forests, estimated to
however, US environmentalist Bill contain 50 percent of the world’s
… the conservation of McKibben declared that calculating species, are particularly vulnerable
natural resources is the a peak oil date was pointless; if all to deforestation. Around 17 percent
fundamental problem. known oil reserves were burned, of the Amazon rain forest alone has
Unless we solve that the carbon produced would be been lost in the past 50 years. As
problem it will avail us five times the amount required “Our Common Future” suggested,
little to solve all others. to heat the planet by 3.6°F (2°C)— part of the problem is that
Theodore Roosevelt the “safe” temperature limit that developing countries can earn
climatologists had worked out in money from large corporations if
2009. The science has evolved, they clear rain forests for mining,
but the predicted risks of burning logging, and cash crops. In
fossil fuels remain dire. Indonesia, for example, intensive
deforestation took place to make
Saving trees way for palm oil plantations.
industrialized nations were on an Forests are a valuable natural asset Greenpeace reports that the amount
economically viable supply. With that Earth cannot afford to lose. of Indonesian rain forest logged,
this came, too, the realization that Their diminished numbers pose a burned, or degraded in the last 50
oil is a finite resource. Scientists significant threat for the climate; years is equivalent in area to twice
had already considered the problem trees are “carbon sinks,” meaning the size of Germany. The UN and
and calculated the date when the they take in carbon dioxide and use other bodies now offer developing
supply of oil would peak, before it it to fuel growth. This then prevents countries technical advice and
ran out or became uneconomical to carbon from contributing to global financial incentives to manage their
extract. In 1974 the peak oil date warming. Trees are a renewable forests in a more sustainable way.
was predicted to be 1995, with the resource, and people, businesses,
caveat that there were several and nations often plant them to Deteriorating soil
potential variables and unknowns offset fossil fuel use, but not in Topsoil is perhaps one of the
such as consumption rates, sufficient numbers. According to world’s most undervalued
available technology, and reserves Friends of the Earth, the annual resources. This vast ecosystem,
Easter Island stone, but logs were needed

as rollers to transport them from
The fate of the ancient people the quarries to ceremonial sites.
of Easter Island illustrates As the island’s many palms were
the importance of managing cut down, there was no wood
natural resources. Once a left for fishing canoes, which
thriving community of 12,000 led to many people starving
people who erected enormous to death.
stone monuments, they had The final tragedy came in
dwindled to just a couple of 1862, with the arrival of slave
thousand by the time Europeans traders, who captured 1,500
discovered the island in 1722. islanders and took them to Peru,
Mismanagement of a fragile where almost all of them died.
ecosystem, especially mass The 15 islanders who eventually
deforestation, and warring managed to make it home Some 887 moai cover the slopes of
between tribes, had been the unwittingly introduced smallpox Rano Raraku, Easter Island’s volcanic
cause of their demise. The giant to the island. By 1877, only 111 crater, the source of the stones from
heads, or moai, are made of inhabitants survived. which the statues were carved.
Thick forests like the one in this locked away in glaciers or in deep innovative ways to minimize
15th-century painting by Italian artist underground aquifers. Only one- environmental impacts and
Paolo Uccello are returning to Europe, hundredth of 1 percent of all the manage resources.
where they have grown by 42 million
acres (17 million ha) since the 1990s.
water in the world is readily Some progress is being made,
available for human use. Drinking in part thanks to campaigning by
water is also not distributed people like Naomi Klein. A number
composed of animals, microbes, equally, being naturally scarcer in of European and Asian countries,
plant roots, and minerals, is a hot, arid areas of the world than including the UK, have decided
complex and delicate structure that in temperate zones. to phase out fossil-fuel vehicles.
is slow to form and easily lost. The Population pressures and wealth In other areas, however, socio-
World Wildlife Fund estimates that also have an impact on water economic and political problems
half of the world’s topsoil has been supplies. The UN believes everyone remain obstacles to reform. As “Our
eroded by wind and rain in the last should have access to at least 88 Common Future” stated, meeting
150 years. Particles then collect in pints (50 liters) of freshwater a day, humanity’s goals and aspirations
streams and rivers, clogging them but people in sub-Saharan Africa responsibly “will require the active
with sediment. Soil loss occurs manage on 21 pints (10 liters) a day, support of us all.” ■
due to overgrazing, removal of while the average American enjoys
hedges, and use of agrochemicals almost 740 pints (350 liters).
that affect the soil structure. Around the world, water sources
Meaures such as resting the soil, are also being bought up by large
terracing, dams, and strategic corporations. Some scientists warn
planting can help. In the village of that, if our current usage patterns You have to
Aamdanda in Nepal, for example, continue and population rates grow think in terms of the
steep-sided slopes are stabilized at their current rate, by 2030 global survival of human
with broom grass. The plant binds demand for clean water will exceed society … it is not only
the soil; it is also a fodder crop supply by 40 percent. the magnitude of
and is used to make brooms,
change, it’s the pace
which the villagers sell. Future plans
New strategies are evidently
at which it changes.
Water pressures required to save the world from
Benjamin Horton
British geographer
Clean drinking water is a limited human destruction. Transition
resource. Water covers around 75 engineering, an emerging multi-
percent of Earth’s surface, but disciplinary field, may help. It
97.5 percent of it is salt water. Of aims to use existing businesses,
the remaining 2.5 percent, most is organizations, and systems to find
266
IN CONTEXT
NOW YOU’VE GOT KEY FIGURE

John Crosbie (1931–)
BIGGER AND BIGGER

BEFORE
1946 The International
Whaling Commission is set up
BOATS CHASING
to review and control whaling,
reversing a dramatic decline
SMALLER AND
after centuries of hunting.
1972 Overfishing and a strong
El Niño cause Peru’s coastal
FEWER FISH
anchovy fisheries to crash—a
blow to the national economy.
AFTER
OVERFISHING 2000 The World Wildlife Fund
places cod on its endangered
species list and launches a UK
Oceans Recovery Campaign.
2001 Jeremy Jackson and
other marine biologists trace
the history of overfishing.
2010 The UNESCO Aichi
Biodiversity Target 11 calls for
a tenth of coastal and marine
areas to be protected by 2020.
I
n 1992, one piece of legislation
changed the ecological,
socioeconomic, and cultural
structure of Canada’s Atlantic
Maritime provinces. John Crosbie,
the Federal Minister of Fisheries
and Oceans, placed a moratorium
on the Atlantic cod fishery; no
further cod could be harvested
from the ocean. His ruling was a
necessity; the volume of northern
cod was down to 1 percent of
previous levels. The region had
been overfished to the point where
recovery could not occur if fishing
were allowed to continue. Crosbie
called it the toughest political
moment of his career. The decision
put thousands of Canadians out of

See also: A holistic view of Earth 210–211 ■ Pollution 230–235 ■ Human
devastation of Earth 299 ■ Sustainable Biosphere Initiative 322–323
Marine reserves
A promising tool for fish
management is the creation of
marine protected areas
Trawlers catch too many fish (MPAs), which legally protect
fish stocks and ecosystems.
MPAs cover around 3.5
percent of the world’s oceans,
but only 1.6 percent of MPAs
are the strongest “no-take
Large predator zones” where fishing,
Immature fish Mature fish fish extraction of materials,
Do not live to Loss of breeding Changes balance dumping, drilling, and
reproduce stock of species in dredging are banned. One
ecosystem meta-analysis of scientific
studies showed that the
volume of fish species is on
average 670 percent greater
in fully protected “no-take”
marine reserves than in areas
Fish stocks decline or become unsustainable that have no protection, and
343 percent greater than in
partially protected MPAs.
No-take zones effectively
preserve and restore damaged
ecosystems, too; coral reefs in
Solution: protected zones of the Pacific
Solution: Solution:
Set quotas and/ Line Islands recovered from an
Set minimum Institute lengthy
or institute El Niño event within a decade,
size limits moratoriums
moratoriums but those in unprotected areas
did not. Some studies suggest
that legally enforced reserves
may even help replenish
fisheries outside their borders.
jobs. For 500 years, the cod fishery conditions in which both the cod
had supported Maritime residents, and its food sources struggle to
particularly in Newfoundland. survive. A further blow to
The 1992 moratorium was Newfoundland’s fishermen—who
initially supposed to last only two largely turned to catching shrimp
years, but, with the stocks not yet and crab—is that where cod
recovered, it is still largely in place. numbers improved, cod began
From around 2005 to 2015, the eating the shrimp. The ecosystem
volume of northern cod rose by cannot support both a large-scale
about 30 percent each year along shrimp and crustacean industry,
Newfoundland’s northeast coast, and large-scale cod fishing.
although stocks further south did
not recover as fast. In 2017 and A sustainable harvest
2018, however, cod numbers The Newfoundland problem
declined sharply, and the overall demonstrates the complexities of Bigeye trevallies are among the
stocks are still too low to support fishery management, which often many species in the Malpelo Fauna
and Flora Sanctuary, the biggest
large-scale fishing. Climate change relies on the concept of maximum no-fishing zone in the Eastern
has contributed to the problem: sustainable yield: the volume of fish Tropical Pacific, noted for its sharks.
higher temperatures have created harvested from the sea should be ❯❯
268 OVERFISHING AND THE GLOBAL MARINE FISHERIES CRISIS

Disrupting the ecosystem
Large-scale fishing
operations disturb the balance
of marine ecosystems in
various ways, depleting the
target fish species, upsetting I didn’t take
the food chain, and damaging the fish from the
the marine environment. goddamn water.
John Crosbie
Bycatch
Physical impact
of fishing gear
Harvest Incidental demand.

mortality mortality The management strategies
Habitat
adopted depend on the nature of
modification the problem. If fish are being taken
Discarded
or destruction bycatch before they are mature, this will
De c
and offal limit the stock’s future ability to
line i reproduce at a maximum level and
n me a n
trophic level
keep their numbers replenished.
Placing minimum size limits on
Biological
interactions fish can help control this type of
overfishing. If too many mature fish
are being caught, this could leave
too few to reproduce and replenish
Altered ecosystem structure and function the present population. In this case,
moratoriums and quotas are among
measures that can help. Finally,
ecosystem overfishing occurs when
equal to the volume replenished short-term economic gains of a fishery is so depleted that the
through reproduction. This is catching more fish rather than on ecosystem itself changes and is
usually achieved through quotas, long-term sustainability. Fishery no longer able to support the fish
which limit the number of fish that management can be further stock at a sustainable level. It
can be brought in during a season. complicated by factors such as the generally occurs when large
Quotas can curb unsustainable open access nature of the ocean, predatory fish are overfished,
fishing: for example, 16 percent of illegal fishing, and an absence of allowing populations of smaller
fish stocks in American waters regulation and oversight. forage fish to increase and alter the
were overfished in 2015, down from entire ecosystem. This happened
25 percent in 2000. However, the A worldwide crisis in the North Atlantic cod fishery:
quota system can encourage Overfishing is now a global issue, without the cod to keep them in
fishermen to take the largest fish with more than 30 percent of the check, the cod’s three main food
possible, and to throw back smaller world’s fisheries harvested beyond sources—shrimp, crab and capelin
fish, which frequently die from the their biological limits, and 90 fish—all increased in numbers.
stress of being caught. In many percent of fish stocks currently The overfishing problem is now
cases, quotas are also not set at a at their limits or overfished. compounded by climate change
truly sustainable limit; commercial Sustainable management is now and pollution, which are also
fishermen often have considerable essential if fisheries are to continue affecting ocean ecosystems. The
lobbying power, and focus on the to provide jobs and meet consumer consequences could be dire. If

global warming continues, it will annually in the 1960s to more than Effects of pollution
cause higher ocean temperatures, 44 lb (20 kg) in 2016. Global demand
sea ice will melt further, and wind is predicted to reach around 236 Two main types of pollution
damage marine ecosystems.
and ocean current patterns will million tons by 2030. Aquaculture,
Runoff from fertilizers is a
change. As a result, nutrients from the farming of fish and seafood, has common problem: the nitrogen
the upper ocean will be transferred begun to meet much of the demand and phosphorus that many
to the deep ocean, starving marine and has the potential to reduce the contain produce algal blooms
ecosystems and reducing pressures on wild fish stocks. (overgrowths of algae, or
photosynthesis by phytoplankton, However, aquaculture has its own phytoplankton), which later
which serve as the base food in the problems. Nutrients and solids die. As they decompose, they
ocean food chain. Within three added to the water can cause the take up oxygen, creating a
centuries—by 2300—the world’s environment to degrade. The “dead zone” in the water that
fisheries could be 20 percent less buildup of organic matter from cannot sustain life. Because
productive, and between 50 to 60 many fish in a farm can change the fish must leave such water or
percent less productive in the North sediment chemistry, which has an perish, juvenile fish living
Atlantic and western Pacific. The impact on the surrounding water. close to the shore are at risk
predictions, calculated in 2018 by Fish may escape, introducing alien before they move into the open
ocean. In 2017, the annual
scientists at the University of species or diseases into the outside
dead zone in the Gulf of
California, Irvine, are based on freshwater or marine environment.
Mexico was more than
extreme global warming—a 17°F While fish farming helps meet 8,500 sq miles (22,000 sq km).
(9.6°C) increase, but their models demand, overfishing still poses Plastic pollution is another
show that it is a possibility. huge dangers for the health of the threat because fish eat it and
world’s marine ecosystems, and the get caught in nets and debris.
Finding new solutions economic future of many nations. Estimates suggest there are
Seafood consumption has risen The Canadian moratorium severely more than 5 trillion pieces of
from 21.8 lb (9.9 kg) per capita disrupted the economy and culture plastic in the ocean, with
of Newfoundland and neighboring over 8 million tons added
maritime provinces. To avoid such each year. If plastic pollution
A deep-sea salmon farm, built in continues unchecked, the
China, begins its journey to Norway. crises, more governments will have
to develop sustainable fishing volume of plastic in the
The huge, semi-submersible cylinder
practices, and protect the health ocean will exceed that of
aqua-farming platform is designed to
produce 1.5 million salmon a year. of ecosystems and fish stocks. ■ fish by 2050.
Thick blooms of phytoplankton

appear in red on this satellite image
of the Gulf of Mexico. Bacteria break
down decaying algae, releasing CO2
and absorbing essential oxygen.
270
IN CONTEXT
THE INTRODUCTION KEY FIGURE

Ryan M. Keane, Michael J.
OF A FEW RABBITS
Crawley (1949–)
BEFORE
1951 The International Plant
COULD DO
Protection Convention is set up
to prevent the introduction and
LITTLE HARM
spread of pests of plants and
plant products as a result of
international trade. It is
adopted in many countries.
INVASIVE SPECIES 1958 The Ecology of Invasions
by Plants and Animals by
British ecologist Charles
Elton is the first book to be
published on invasion biology.
AFTER
2014 Studies of some of
the “world’s worst” invasive
species by ecologists at
Queen’s University, Belfast,
and Stellenbosch University,
South Africa, reveal that the
ecological impacts of these
species could be predicted
from their behavior.
S
ome of the greatest damage
to ecosystems is caused by
invasive species. These are
plants, animals, or other organisms
that are not native to an ecosystem
but introduced largely through
human action, either deliberately
or by accident. They can become
competitors, predators, parasites,
and hybridizers of native plants and
animals, ultimately threatening the
survival of those species.
The rise of the rabbit

One of the most notable species
invasions has been that of the
European rabbit in Australia.
It began in 1788, when 11 ships

See also: Predator–prey equations 44–49 ■ Non-consumptive effects of predators on their prey 76–77 ■ Human activity and
biodiversity 92–95 ■ The food chain 132–133 ■ The ecosystem 134–137 ■ Chaotic population change 184
Spread of rabbits in Australia
The harlequin ladybug is the world’s

most invasive ladybug. In the UK,
where it was first seen in 2004, it
is reportedly responsible for the decline
of seven native ladybug species.
landed at Botany Bay from Britain,

to establish the first Australian
penal colony. On board the “First
Fleet,” along with more than 1,000 Since their arrival
people, including convicts and in Australia, rabbits
emigrants, were six European have spread throughout KEY
rabbits, brought along for food. the country. They have
< 1870 1900–1910
By the 1840s, rabbits had contributed to the decline
become a staple food in Australia, of many native plants and 1870–1880 1910–1920
and were contained within stone animals, and may have 1880–1890 > 1920
caused several small
enclosures. All this changed mammals to go extinct. 1890–1900 Not included in analysis
in 1859 when a settler, Thomas
Austin, imported 12 pairs of
European rabbits and released There have been several attempts without accidentally introducing
them on his estate near Geelong in to control the feral population, from additional ecosystem problems.
Victoria. Twenty years later, rabbits rabbit-proof fences stretching more Despite being hampered by the
had migrated to South Australia than 2,000 miles (3,200 km) to the lack of comparative data on those
and Queensland, and then in the more successful introduction of the invasive species that fail to
next two decades to Western myxoma virus and the rabbit succeed, scientists have developed
Australia. By 1920, the rabbit calicivirus, in 1950 and 1995 a number of theories to explain the
population was 10 billion. respectively. The resulting disease success of certain species in
Rabbits appear to be innocuous has proved the most effective way nonnative environments, including
creatures, but they have wreaked of controlling their numbers and the resource availability hypothesis,
havoc on Australia’s native species, protecting native species. the evolution of increased
competing with them for resources competitive ability hypothesis, and
such as grass, herbs, roots, and The secrets of success the enemy release hypothesis.
seeds, and degrading the land. As invasive species have spread In general, species success
They become particularly throughout the world, scientists depends on a variety of genetic,
troublesome during a drought, have tried to determine what ecological, and demographic
when they eat anything they makes some of these species so factors. The resource availability
can find in order to stay alive. successful, and how to control them hypothesis, first proposed in 1985 ❯❯
272 INVASIVE SPECIES

The zebra mussel
The case of the zebra mussel They have fewer They are well
demonstrates the diverse enemies than suited to the new
ways to approach invasive native species. environment.
species control, and the
challenges that result. Zebra
mussels are small, fingernail-
sized mollusks with a dark-
striped shell. The mussel is
Why invasive species succeed
native to Eurasia but was
discovered in the Great Lakes
area of North America in 1988,
probably carried there in
ballast water discharged from They out-compete They are toxic
ships traveling from Europe. native species. to native species.
Since then, zebra mussels
have spread throughout the
midwestern United States,
and have been found as far
by the ecologists Phyllis Coley, mustard (Alliara petiolata). Native
west as California.
The zebra mussels attach John Bryant, and F. Stuart Chapin, to Europe, western and central
themselves to clams and other argues that an invasive species Asia, and northwestern Africa,
mussels, filtering out algae thrives because it is already well it was brought to North America
that the native species need suited to its new environment by early settlers to use in cooking
for food to survive. They also and can take advantage of any and medicines, and rapidly spread.
clog water intake pipes used surpluses in resources. The Continued infestation has affected
for power plants and drinking evolution of increased competitive the growth rate of tree seedlings
water supplies. Current ability hypothesis, published by and reduced the native plant
control mechanisms include ecologists Bernd Blossey and diversity, leading to changes in
chemicals, hot water, and Rolf Nötzold in 1995, suggests the forest ecosystems invaded.
filtering systems. While each that invasive plants facing fewer In its native range, garlic
has had some success, none herbivores in their naturalized mustard is consumed by as many
of these solutions has been environment can allocate more as 69 insect species, but none of
capable of safely eradicating
resources to reproduction and these is present in North America.
the mussels. As a result, they
continue to spread throughout
survival and so out-compete the This lack of predation and the
the waterways of the US. native species. The enemy release plant’s invasive success provide
hypothesis, set out by ecologists
Ryan M. Keane and Michael J.
Crawley in their 2002 article “Exotic
Plant Invasions and the Enemy
Release Hypothesis,” argues that the
invasive species has fewer enemies
in its naturalized environments, and
We are seeing one of the great so can spread farther. The reality is
historical convulsions in the that the success of invasive species
world’s fauna and flora. is likely due to many mechanisms
Charles Elton working together.
Garlic mustard is highly invasive
Plant invaders in North America, inhibiting other
One plant that appears to support plants. In its native habitat, it is
multiple hypotheses about the considered an attractive wildflower,
success of invasive species is garlic although it can have a strong smell.

Since their introduction to
Australia in 1935, cane toads have
out-competed native frogs because
they reproduce far more quickly.
support for the enemy release

hypothesis. Garlic mustard also
successfully competes with native
plants for resources, fulfilling the
resource availability hypothesis.
The plant even exudes secondary
compounds that may “attack”
native plants by inhibiting their
germination and growth. This
supports the “novel weapons”
hypothesis, proposed by ecologists
Wendy M. Ridenour and Ragan M.
Callaway in 2004, which posits that
invasive species have biochemical
weapons that give them a key
advantage over native species.
The art of control

Successful invasive species are Not all biological controls are the economy of local communities.
extremely difficult to control and effective, and some measures have Regulators are, therefore, often
almost impossible to eradicate. had disastrous consequences. For hesitant to support biological
If the species is a plant, the most example, in 1935 cane toads were controls without extensive prior
obvious way to remove it is to introduced to Australia to control research. No magic bullet exists
pull it up or cut it down, but such the invasive grayback cane beetle, that can control every invasive
methods are highly labor-intensive, which was destroying sugar cane species. They are dependent on
especially over a wide area. The fields. The cane toad had been complex ecosystem interactions,
use of chemicals to destroy effective in controlling beetles and scientists continue to design
invasive species is often in Hawaii, so the assumption was field experiments to test their
successful, but it can also kill that it would be equally successful hypotheses of how invasive
native species and undermine soil in Australia. However, grayback species function in the wild. ■
health, with the added threat of cane beetles feed primarily at the
harm to humans. top of sugar cane stalks, which is
One frequently used method out of reach for the cane toads.
of control, known as biological A lack of understanding of the
control, or “biocontrol,” pits an different environments favored
invasive species’ own enemies by the two creatures meant that Now is the time to take action.
against it. In an early success, the cane toad was the wrong
The costs to habitats and the
the cactus moth was introduced choice as a biological control. By
to Australia from South America the time the mistake was realized,
economy are … out of control.
in 1926 to feed on the prickly the toad had spread throughout
Bruce Babbitt
US Secretary of the Interior ,
pear. This plant had itself been Australia, poisoning any predator (1993–2001)
introduced in the 1770s and was species that tried to eat the
choking farmland in New South toxic amphibian.
Wales and Queensland. By the Even when biological controls
early 1930s, most prickly pears curb an invasive species, they may
had been eradicated. create imbalances in ecosystems or
AS TEMPERATURES INCREASE,
THE DELICATELY
BALANCED SYSTEM
FALLS INTO
DISARRAY
SPRING CREEP
276 SPRING CREEP
M
ost scientists now agree may be triggered by temperature,
IN CONTEXT that climate change, rainfall, or the length of daylight,
driven by an increase but temperature is probably the
KEY FIGURE
in greenhouse gases, is raising single most important factor in
Camille Parmesan (1961–)
the global mean temperature. The Earth’s temperate and polar regions,
BEFORE Intergovernmental Panel on Climate whereas rainfall is the key factor in
1997 A group of American Change (IPCC) cites an increase the tropics. In 2003, climate change
scientists publishes evidence of 1.8°F (1°C) since 1880, although scientists Camille Parmesan and
of a longer plant growing in some regions the warming has Gary Yohe proved that spring
season at northern high been even more marked. This change is now happening earlier—
latitudes in 1981–91. warming has affected both plant a phenomenon called spring creep.
and animal behavior, and the IPCC
2002 Naturalist Richard Fitter forecasts a further increase Season creep
reveals that the first flowering of 2.5–9.9 °F (1.4–5.5°C) during the For several decades, people have
date of 385 plant species in the next 100 years. observed leaves and flowers
UK has advanced by 4.5 days The life cycles of plants and appearing earlier in spring. In
in the previous decade. animals change in line with the the past, these claims were often
seasons. Phenology is the study dismissed as lacking “hard science,”
AFTER of these seasonal changes. They such as facts, figures, or datasets.
2006 Jonathan Banks, from
the American Clean Air Task
Force, is the first person to use The impact of seasonal
the term “season creep” to changes on plants and animals
describe the increasingly early
onset of the seasons as a result
of climate change.
2014 In the US, the National
Climate Assessment confirms Plants grow leaves, Mammals breed and raise
long-term trends toward produce flowers and fruit, young. Some mammals go into
shorter, milder winters and and shed their leaves. hibernation over the winter.
earlier spring thaws.
Seasonal changes
in the weather
We are seeing
change happen
much faster than
I thought it would Birds nest and breed. Many birds After hatching, amphibians, insects, and
10 years ago. (and some other animals) make some other animals metamorphose from
Camille Parmesan long-distance migrations. one body form into another.
All life forms respond to changes in weather brought

about by the seasonal cycle. Migration, breeding,
flowering, hibernation, and metamorphosis are
some of the events affected by this cycle.

See also: Animal ecology 106–113 ■ Animal behavior 116–117 ■ The foundations of plant ecology 167 ■ Global warming
202–203 ■ Endangered habitats 236–239 ■ Halting climate change 316–321
When Camille Parmesan and Gary than before, providing a longer

Yohe published evidence in 2003— growing season for plants. As
based on an analysis of more than some regions become drier and
1,700 species—they demonstrated some wetter, bouts of extremely
that change was very real. Their heavy rainfall and flooding have
data showed that spring change become more common. Toxic algal
was indeed taking place earlier—by blooms in lakes are occurring more
an average of 2.3 days per decade. frequently. Ice cover in polar regions
Studies by other scientists in recent is also decreasing. All these changes
years have supported their findings. have affected and will continue to
Many of the changes that take affect animal and plant behavior.
place in plants are governed by Since 1993, the European
temperature, including growth Environmental Agency (EEA) has
spurts; the appearance of leaves, worked in earnest to pull together
flowers, and fruit; and leaves dying data from thousands of studies—
in fall. Most food chains start with dating back to at least 1943—to
plants, so these changes affect create a picture of spring creep in The leaves of some oak species
grazers and browsers, from rabbits Europe. The EEA’s evidence shows turn red shortly before they fall in fall.
to deer, and pollinators, including earlier dates for plants producing Comparing the date on which this
occurs from year to year can provide
bees and butterflies. All of these pollen, frogs spawning, and birds evidence for climate change.
are at the bottom of the food chain nesting. According to their data,
(primary consumers). If they many insects whose life cycles
struggle to find food, those that prey are governed by air temperature In Spain, botanists studied data for
on them (secondary consumers) also (thermophilic insects, such as 29 species of plants. They found
suffer from the absence of prey. butterflies and bark beetles) now that in 2003, leaves first appeared
have a longer breeding season, 4.8 days earlier on average than
Effects of climate change enabling them to produce extra in 1943; flowers first bloomed
A warmer Earth produces many generations each year. For example, 5.9 days earlier; trees produced
effects. In most cooler parts of the some butterflies that previously had fruit 3.2 days earlier; and leaves
world, the frost-free season is longer two generations now have three. died 1.2 days later. In the UK, the ❯❯
Camille Parmesan Born in 1961, Professor Camille figure in the IPCC and her work
Parmesan is an American has won her many accolades
academic who has established and has been cited in hundreds
a reputation as one of the leading of academic papers. She is
climate change scientists. She professor in Integrative Biology
received her Ph.D. in biological at the University of Texas at
sciences from the University of Austin and advises international
Texas at Austin in 1995 and her conservation bodies.
early research concerned the
evolution of insect–plant Key works
interactions. For the best part of
20 years, she has focused on 2003 “A globally coherent
documenting the shifting fingerprint of climate change
geographical ranges of butterflies impacts,” Nature
across North America and Europe, 2015 “Plants and climate
linking these to climate change. change: complexities and
Parmesan has been a leading surprises,” Annals of Botany
278 SPRING CREEP

evidence was even more dramatic: undergo explosive population their destination 25 days sooner
across 53 plant species, leaves, increases and produce damaging than previously. However, there is
flowers, and fruits appeared almost infestations. Warmer springs allow evidence that birds migrating from
six days earlier in 2005 than they pine sawflies, whose larvae eat Central America to New England in
had done in 1976. Similarly, the pine needles, to develop too rapidly the US have declined faster than
fruiting season of 315 different for the birds and parasites that feed birds that remain in New England
kinds of fungi studied in Britain on them to keep their numbers in all year. This is probably because
lengthened from 33 to 75 days in check. Out of control, the sawflies the migrant birds have been unable
the second half of the 20th century. strip trees of their needles and to adjust their departure dates from
Longer plant growing seasons stunt their growth. Central America to arrive in time to
sound like good news, but warmer benefit from the earlier abundance
temperatures create problems as Migration and hibernation of insects the way local birds do.
well as advantages. Not all insects Birds that migrate in spring to Climate change also appears
are welcome, and shorter, milder reach rich food sources also face to have changed the behavior of
winters kill fewer dormant insects, problems. Some have adjusted their hibernating mammals. Zoologists
some of which may consequently flight schedules to benefit from the at the Rocky Mountain Biological
earlier abundance of insects. After Laboratory found that yellow-
making the long journey from sub- bellied marmots living in Colorado
Some bee species now emerge earlier
in spring, in line with earlier flowering Saharan Africa, the first swallows emerged 38 days earlier in 1999
dates for the plants that they pollinate. arrive in the UK about 20 days than they had done in 1975. In
Other bees, though, have not been able earlier than they did in the 1970s, 2012, scientists at the University
to synchronize their emergence. and the first Bank Swallows reach of Alberta found that in the last two
decades, late snowfall has delayed
the emergence of the Rocky
Mountain ground squirrel from
hibernation by 10 days. This has
cut down the already short active
period in which they mate, give
birth, and feed to prepare for the
next hibernation cycle.
Decoupling
Some organisms’ survival could be
threatened by the “decoupling” of
interactions between species. This
We are now sure of what we

only suspected years ago.
Policy needs to catch
up with science.
Camille Parmesan

Wall butterflies and
climate change
Climate change sometimes
produces unexpected results.
For example, in the UK, the life
cycle of the wall butterfly has
been disrupted by changing
climatic conditions. Previously,
the butterfly produced two
generations every summer.
The late-summer adults would
mate, the females lay eggs,
and the eggs then developed
into caterpillars. In September,
these caterpillars found
sufficient food to grow large
and sustain themselves in
hibernation through winter.
In spring, the caterpillars
metamorphosed into pupae,
and then became adults.
A Great Tit feeds its chicks. If a problem for birds that depend
breeding takes place after the peak Warmer weather has allowed
on peaks in insect abundance. For a third generation to develop
period for spring caterpillars, there example, Pied Flycatchers and
will be less food for the young birds, in fall, with adults flying as
and fewer will survive to breed.
Great Tits feed their chicks on late as mid-October. By
caterpillars that are abundant for a the time the third generation
short period in spring. Due to caterpillars hatch there is
could seriously upset the balance climate change, the caterpillar little food, so most starve
of ecosystems. If flowers appear peak is now earlier, but the birds and die. Scientists call this
earlier, the bees that pollinate them have not been able to advance a “developmental trap” and it
can respond in one of two ways: their egg-laying dates enough to is probably responsible for the
they can emerge earlier; or they can take advantage of the glut of food. decline in wall butterflies.
move to a higher latitude to match Studies show that fewer Pied
later flower emergence farther from Flycatcher and Great Tit chicks
the equator. Studies of 10 wild bee are surviving. Pied Flycatcher
species in northeast North America numbers have declined in Dutch
have shown that their behavior woodlands, possibly as a result of
has changed in line with earlier climate change.
flowering. However, bumblebees in
Colorado have not matched the Taking action This butterfly I was
changes and their population has All of this disturbing evidence studying shifted its
fallen. If pollinators decline, so may has prompted climate scientists entire range across
the plants that they pollinate. worldwide to lobby governments half a continent—
There is evidence that many and demand policy change. Spring I said this is big …
primary consumers have adjusted creep has been used by scientists Everything since then
to changed natural phenomena, but as a definitive piece of proof that has just confirmed it.
species higher in the food chain climate change is occurring, and Camille Parmesan
seem to find it harder to make the researchers have called upon policy
change. Although birds are now makers to fight global warming to
nesting earlier than they once did, save the familiar species that find
the timing of insect emergence has their very existence threatened
advanced more rapidly. This is by phenological changes. ■
280
ONE OF THE MAIN

THREATS TO
BIODIVERSITY IS
INFECTIOUS
AMPHIBIAN VIRUSES
DISEASES
S
ince the 1980s, hundreds of
IN CONTEXT species of amphibians have
suffered population crashes
KEY FIGURE
and localized extinctions—at a rate
Malcolm McCallum
thought to be more than 200 times
(1968–)
the natural, “background” extinction
BEFORE rate unaffected by modern human
1989 The formerly common activity. This alarming phenomenon
golden toad of Costa Rica is first attracted public attention in
declared extinct. Various 1999, when American environmental The North American bullfrog is
scientist Malcolm McCallum resistant to the chytrid fungus, but
explanations are proposed. acts as a deadly carrier of the infection
published his findings about the
1998 In the US, many poison- dramatic increase in deformities to other species of amphibians.
dart frogs die at the National in frogs. He went on to produce
Zoo in Washington DC. The landmark studies on amphibian they are not able to breathe, hydrate,
chytrid fungus is implicated decline and extinction. or regulate their temperature. The
as a cause. The causes of the problem are exact origin of the fungus is not
wide-ranging, and include habitat known, but the global trade in live
AFTER destruction and pollution, as well as amphibians for various uses, be it
2009 The Kihansi spray toad competition from nonnative species. pets, food, fishing bait, or research,
of Tanzania is declared extinct But one of the most devastating has been a major factor in its spread.
in the wild as a result of chytrid causes is undoubtedly disease, with Ranaviruses evolved from a fish
infection. two particularly lethal culprits. virus. They infect amphibians and
reptiles, and have caused mass
2013 A second species of
Chytrid and ranavirus mortality in frogs since the 1980s.
chytrid fungus causes the near- Chytridiomycosis is a disease The common midwife toad ranavirus
extinction of fire salamanders caused by the chytrid fungus, and causes bleeding, skin sores, lethargy,
in the Netherlands. it has ravaged populations of frogs and emaciation. It is notably virulent
2015 The chytrid fungus is and toads in particular. The fungus as it has the ability to “jump” from
detected in amphibians in 52 affects amphibians’ skin, such that one species to another. ■
out of 82 countries sampled.
See also: Biomes 206–209 ■ Pollution 230–235 ■ Endangered habitats 236–239
■ Deforestation 254–259 ■ Overfishing 266–269
IMAGINE TRYING TO
BUILD A HOUSE WHILE
SOMEONE KEEPS
STEALING YOUR BRICKS
OCEAN ACIDIFICATION
A
dding carbon dioxide (CO2) of CO2 pollution on the oceans.
IN CONTEXT to the air not only triggers They took samples of seawater
climate change but also from around the world, and found
KEY FIGURES
makes the oceans more acidic. So that the acidity had increased
Kenneth Caldeira (1960–),
far, the oceans have buffered the measurably in the past 200 years
Michael E. Wickett (1971–) worst effects of global warming, of industrialization. They coined
BEFORE absorbing up to half of the carbon the term “ocean acidification” and
1909 Danish chemist Søren dioxide added to the atmosphere by predicted that this change could
Sørensen develops the pH human activity. However, the gas accelerate over the next 50 years,
scale for measuring acidity. alters the oceans’ chemistry. with damaging results.
In 2003, American climate Many sea creatures rely on the
1929 American biologists scientists Ken Caldeira and Michael natural alkalinity of seawater to
Alfred Redfield and Robert E. Wickett investigated the effects maintain carbonates for building
Goodkind discover that excess their shells and skeletons. Even a
carbon dioxide in water slight decrease in alkalinity seriously
suffocates squid. disrupts growth, especially for
sensitive creatures such as corals
1933 German chemist and plankton. Acidification might
Hermann Wattenberg makes wipe out corals within decades; if
the first global survey of ocean Most carbon dioxide released they go, so do the reef ecosystems.
acidity, as he analyzes results into the atmosphere as a result Phytoplankton are the foundation
from the Atlantic expedition of burning fossil fuels will be of the ocean food web, and are
of the Meteor research vessel. absorbed by the ocean. vital to maintaining global
Ken Caldeira and oxygen levels.
AFTER Michael Wickett Ocean acidification is far harder
2012 In the US, oceanographer to reverse than the atmospheric
James C. Zachos and his effects of CO2 emissions, and its
colleagues use fossil evidence devasting impact on biodiversity,
from marine sediments to fisheries, and food security remains
show that past acidification a serious concern. ■
of the ocean has led to mass
extinctions of sea creatures. See also: Global warming 202–203 ■ Pollution 230–235 ■ Endangered habitats
236–239 ■ Acid rain 248–249 ■ Halting climate change 316–321
282
THE ENVIRONMENTAL
DAMAGE OF URBAN
SPRAWL CANNOT
BE IGNORED
URBAN SPRAWL
S
ince the 1950s, the term Modern definitions of urban sprawl
IN CONTEXT “urban sprawl” has been vary, but it generally has negative
widely used to describe the overtones. At its most extreme, it
KEY FIGURE
growth of low-density suburbs has created megacities—defined by
Robert Bruegmann (1948–)
beyond high-density city cores. the United Nations as cities of more
BEFORE The term was first used by The than 10 million people. Examples
1928 British architect Clough Times newspaper in the UK in 1955 of such megacities include Tokyo-
Williams-Ellis compares to describe the spread of London’s Yokohama (38 million), Jakarta
London’s growth to an octopus suburbs. At this time, the British (30 million), and Delhi (25 million).
devouring the countryside. planning authorities were
introducing “green belts” around Ecological upset
1950s With postwar prosperity cities, where new building was Some researchers claim urban
and increased car ownership in almost entirely banned. Green sprawl is the most serious threat
the US, the middle classes leave belts were designed to stop cities to biodiversity from any human
cramped city centers and move from spreading and merging with activity. The new suburbs house
to new, low-density areas in other towns. relatively few people, yet require
the suburbs. extensive and disproportionate
levels of infrastructure, such as
AFTER power and water supplies and
2017 A housing crisis in the transportation networks. As cities
UK prompts calls for the lifting swell, valuable farmland is covered
of restrictions on new building in concrete and natural habitats are
on the greenbelts around major The old city is disrupted or lost entirely. Sprawl can
UK cities. submerged in a also disturb local fauna and flora
far-flung, multicentered, through the introduction of pets
2050 The date by which, mostly low-density, and invasive plants that threaten
according to UN estimates highly heterogeneous indigenous species. Limited public
published in 2014, the urban urban region. transportation in low density
population of the world is Robert Bruegmann areas also means that suburban
set to rise to 6.34 billion out populations tend to be multiple car
of a projected total population owners, which adds to the levels
of 9.7 billion. of air pollution in cities—as do the
wood- and coal-burning stoves of
the poor in outlying shanty towns.

See also: Pollution 230–235 ■ Endangered habitats 236–239 ■ Deforestation 254–259 ■ Depletion of natural resources
262–265 ■ Amphibian viruses 280
Toluca was once a picturesque old

town to the west of Mexico City. Now
a city of more than 800,000 people, it is
gradually merging into the sprawl of
Mexico City—at a high ecological cost.
The area of the world currently

covered in urban development
is one-and-a-half times the size
of France. Mexico City has expanded
more than any other city in the West.
Spreading far beyond its official
boundaries to become the home
of more than 21 million people, it
has also grown disproportionately:
in 1970–2000, the surface area of the
city grew 1.5 times faster than its
population. While 59 percent of the
city’s territory is conservation land,
illegal logging and urban sprawl
continue to degrade urban forest, blocks, pushing the city limits— million and that of Beijing to 23
grassland, and water supplies. and the urban poor—far from city million by restricting land available
It is estimated that 37 percent centers. The reliance on cars in the for building and controlling the
of all urban growth by 2050 will new neighborhoods, and the lack of inflow of people, forcing out
occur in China, India, and Nigeria central hubs, means there is little low-skilled workers. China is
alone. In Beijing and other cities in opportunity for community life. also building higher-density
China, densely populated hutongs Aware of the problems caused neighborhoods with narrower
(alleyways), where the urban poor by urbanization, the Chinese streets, more intersections, and
used to live, are being demolished government is now trying to limit more public transport that will help
to make way for low-density luxury the population of Shanghai to 25 the formation of communities. ■
The endangered axolotl One of the victims of the urban Historically, the wild axolotl
sprawl of Mexico City has been lived in the urban canals created
the tiny axolotl, a pale-colored by the Aztecs as they built their
salamander that looks like a fish capital city in the 13th century,
but is actually an amphibian, and in the network of lakes
and is sometimes known as the around the city that fed these
Mexican walking fish. Capable of canals. As Mexico City has
growing up to 1 ft (30 cm) long, the expanded, these canals have
axolotl feeds on aquatic insects, been lost, and the wild axolotl
small fish, and crustaceans, and has declined. In 2006, it was
has the ability to regenerate added to the list of critically
severed limbs—a quality that endangered species and by
has made captive specimens an 2015 it was thought that the
important subject of scientific creature may have been extinct.
research. The captive version is However, specimens have since
also a familiar pet in aquariums been found in Lake Xochimilco
around the world. in southern Mexico City.
284
OUR OCEANS ARE

TURNING INTO A
PLASTIC SOUP
A PLASTIC WASTELAND
W
hen plastics were first Charles Moore, who highlighted it in
IN CONTEXT mass produced in the his 2011 book Plastic Ocean. Sailing
early 20th century, the home from a yachting competition,
KEY FIGURE
world marveled at the versatility and Moore came across a vast patch of
Charles J. Moore (1947–)
durability of a material that could be plastic debris in the Pacific Ocean.
BEFORE molded into any shape, used, and Now known to have a bigger surface
1970s Scientists begin to then thrown away. The problem with area than France, Germany, and
research plastic litter at sea plastic, however, is that most of Spain combined, the Great Pacific
after reports in the journal it never goes away. According to Ocean Garbage Patch (GPOGP)
Science describe large the British business publication comprises 79,000 tons of
numbers of plastic pellets The Economist, only 20 percent microplastics amassed by the
in the North Atlantic. of the 6.3 billion tons of plastic swirling current known as the
produced in the world since the North Pacific Gyre.
1984 The first International 1950s has been burned or recycled. The GPOGP is one of several
Marine Debris Conference, This means that 80 percent— oceanic garbage patches—there
held in Hawaii, raises 5 billion tons—is in landfills or are others in the Atlantic and
awareness of the growing elsewhere in the environment.
problem of litter in the oceans.
Polluting the oceans
AFTER Microplastics—tiny fragments of
2016 The documentary A plastic less than 1 ⁄4 inch (5 mm)
Plastic Ocean, directed by across—are even harder to clean
Australian journalist Craig up than other plastics. Comprising
Leeson, highlights the global 90 percent of the plastics in the
effects of plastic pollution. oceans, they surge through currents
like a murky soup. The problem
2018 The Earth Day Network, was first identified in 1997 by the
an organization committed to American oceanographer Captain
spreading the environmental
movement worldwide, makes
End Plastic Pollution the theme A “seabin” is emptied in Sydney
harbor. The Seabin Project, introduced
of Earth Day, on April 22, 2018. in Australia in 2015, helps counteract
plastic pollution by filtering surface
water in ports and harbors.

See also: The food chain 132–133 Humankind’s dominance over nature 296
■ Human devastation of Earth 299
■
■ Man and the Biosphere programme 310–311

Effects on wildlife
Plastics pose a danger to
wildlife in many ways. Larger
items such as plastic shopping
Plastic is recycled, burned, put into bags can choke or strangle
landfill, or dumped in the ocean. birds and marine animals; if
ingested, they can damage
their digestive tracts or cause
starvation by obstructing the
stomach. If microplastics are
It takes thousands of years to decompose. ingested, toxins can pass
into an animal’s fatty tissues,
a process that then passes up
the foodchain.
According to Greenpeace,
Wave action and UV sunlight breaks nine out of ten seabirds, one
plastic in the ocean into tiny fragments, which spread in three sea turtles, and more
through the water. than half the population of
whales and dolphins have
eaten plastic. Even some of
the crustaceans living in the
western Pacific’s Mariana
Our oceans are Trench, the deepest point in
turning into a plastic soup. the world’s oceans, are known
to have ingested plastic.
Companies are starting
to take the need to reduce
Indian Ocean as well as in smaller and many countries, following the plastic use seriously. A brewer
bodies of water such as the North lead of Bangladesh in 2002, are in Florida, for instance, has
Sea. Plastic microbeads, introduced banning the provision of single-use found a way to make six-pack
by cosmetic companies in the plastic bags. Other measures rings from by-products of the
1990s, add to the problem. Used include banning plastic straws brewing process, so that
in personal care products such and promoting the use of reusable seabirds can chew them off
if they become caught in them.
as soaps, facial scrubs, and water bottles and recyclable or
toothpastes, the beads travel from compostable packaging. ■
wastewater systems into rivers and
oceans, where they are consumed
by fish and other animals, with
the same damaging effects as
microplastics (see panel, far right).
Steps to limit plastic The throwaway society

Cleaning up plastic pollution is a cannot be contained—it
gargantuan task. Breaking plastics has gone global. We cannot
down into their constituent store and maintain or
chemicals requires huge amounts recycle all our stuff.
of energy, which also damages the Charles J. Moore
environment. The best solution is A Northern Gannet is entangled
to learn to live without plastic. in the plastic rings of a six-pack.
Birds that scavenge along the shore
Most countries have banned or are such as seagulls are especially prone
working toward phasing out the use to being caught in such debris.
of microbeads in beauty products,
WATER
IS A PUBLIC
TRUST, AND A
HUMAN RIGHT
THE WATER CRISIS
288 THE WATER CRISIS
IN CONTEXT
KEY FIGURE
Maude Barlow (1947–)
BEFORE
1983–5 Droughts in Ethiopia,
Eritrea, and Sudan cause
450,000 deaths.
1990 The desiccation of the
Aral Sea is declared the world’s
worst ecological disaster of
the 20th century by the UN
Environment Programme.
2008 The United Nations
estimates that around 42,000
people die every week from
diseases related to bad water
and poor sanitation.
AFTER
I
2011–17 California suffers one
n 2008, Canadian activist Indians line up for water in a slum
of its worst droughts on record. area of Hyderabad in 2007. India
Maude Barlow argued that
It impacts on agriculture, water shortage had become the suffered a severe water crisis in 2018,
nature, and daily life. most pressing ecological and human
and demand is projected to be twice
the available supply by 2030.
2017 Water campaigner crisis of the 21st century. Stressing
Maude Barlow reveals that that water is a “Commons” (a shared
half of China’s rivers have resource) and that access to water is percent of the planet’s water is both
disappeared since 1990. a fundamental human right, she set fresh and easily accessible. It is
out how wastage, pollution, and obtained mostly from rivers, lakes,
overconsumption meant that the and underground aquifers (rock
water cycle—the constant exchange containing groundwater). People
of water between Earth’s surface use water to drink, wash, irrigate
and the atmosphere—could not crops, and run industry, and since
be relied upon to provide water for all plants and terrestrial animals
evermore. She said that shortage require freshwater to live, all are
of water was already a crisis in the affected by the water crisis.
Life requires access to developing world, where the burden
clean water; to deny the is borne particularly by women and Wasted water
right to water is to deny children who collect water—and A larger human population uses
the right to life. The fight unless drastic action is taken, the more water, and a large proportion
for the right to water is an rest of the world will be affected too. of that is wasted, especially in
idea whose time has come. About 1.1 billion people lack developed countries, where people
Maude Barlow easy access to water, and 2.7 billion on average use about 10 times more
find water scarce for at least one than those in the developing world.
month of the year. Although 70 Sources of freshwater have dried up
percent of Earth’s surface is (for example, much of the Rio
covered by water, almost all of it is Grande between Mexico and the
saline ocean water. Only 0.014 US) or are becoming too polluted to

See also: The ecosystem 134–137 ■ Pollution 230–235 ■ Acid rain 248–249 ■ Overpopulation 250–251
■ Depletion of natural resources 262–265
Distribution of the world’s water

1% Rivers
Easily accessible
1% Water within
freshwater is a very 8% Atmospheric living organisms
fragile resource. Only water vapor
a tiny fraction of the 38% Soil
total amount of water moisture
available on our planet
is immediately fit for 1% Easily accessible
human consumption. 2.5% Freshwater 20% surface water 52% Lakes
Groundwater
97.5% 79%
Oceans Ice caps and
glaciers
All water Freshwater Easily accessible

surface water
use. The Ganges in India and the (64 billion cubic meters)—due in In the last century, half of Earth’s
Citarum in Indonesia are two of the part to the population growing by wetlands have disappeared to make
most polluted rivers in the world. 80 million people each year. The rise way for farmland or development,
At the current rate of consumption in demand has also been driven by or because groundwater has been
this situation will deteriorate further. changing lifestyles and eating removed from aquifers faster than
By 2030, two-thirds of the world’s habits that require more water it has been replaced. A reduction in
population may face shortages. per person. The production of wetlands means plants and animals
Ecosystems will also suffer. biofuels has also risen sharply, with dependent on them are also gone.
significant impact on water Nearly half of all drinking water
Increased demand demand. Between 260 and 1,060 comes from aquifers. About 240
Human use of freshwater has tripled gallons (1,000– 4,000 liters) of water cubic miles (1,000 cubic km) is
since about 1970, and demand is are needed to make about taken every year. Two-thirds is used
increasing by 2,260 billion cubic feet ¼ gallon (1 liter) of biofuel. for irrigation, 22 percent for ❯❯
Maude Barlow Born in Toronto, Canada, in 1947, Peace” nominated for the 2005
Maude Barlow is an activist and Nobel Peace Prize. In 2008, she
water policy critic. She is the received the Citation of Lifetime
author or coauthor of 18 books, Achievement, Canada’s highest
including the bestseller Blue Gold: honor for environmentalism.
The Fight to Stop the Corporate
Theft of the World’s Water. Barlow Key works
formerly served as an adviser
on water to the United Nations, 2002 Blue Gold: The Fight to
and led moves to have water Stop the Corporate Theft of the
recognized as a basic human World’s Water
right. In 2012, she helped found 2007 Blue Covenant: The Global
the Blue Planet Project, which Water Crisis and the Coming
campaigns for the right to water. Battle for the Right to Water
Barlow chairs the Council of 2014 Blue Future: Protecting
Canadians social action group, and Water for People and the
was one of the “1000 Women for Planet Forever
290 THE WATER CRISIS
The desiccation of the Aral Sea

The disappearance of most of the smaller lakes, and contained only
Aral Sea, once the world’s fourth- one-tenth of its 1960 volume of
largest lake, in Kazahkstan has water. Large areas are now
been a huge ecological disaster. desert. Most of the lake’s fish
In the early 1960s, the two main and other aquatic life disappeared
rivers that fed the lake were with its water. Once fishermen
diverted to irrigate millions of here could catch Syr Darya
cotton plants across central Asia. sturgeon, but its numbers
In June 2004, the UN warned that declined sharply when the lake
A stranded ship on the dried-up the lake could dry up completely shrank and became more saline.
bed of the Aral Sea. The loss of such unless measures were taken to Efforts to replenish the waters
a large body of water has had a save it. It was then receiving only have achieved an increase in
devastating effect on agriculture, 10 percent of the water that it surface area and depth, and fish
climate, and the local fishing industry. once did, had divided into several populations are now increasing.
domestic use, and 11 percent for Ogallala aquifer is being depleted. not have plentiful water, such as
industry. However, most aquifers There are even water supply North Africa, the Arabian Peninsula,
replenish much more slowly than problems in Brazil, which is the large areas of central and south
they can be emptied, so water most water-rich nation on Earth. As Asia, northern China, and the
yields reduce with use. If the water the situation worsens, it becomes southwest United States. In contrast,
table falls, some lakes and rivers dry a growing source of conflict. economic water scarcity occurs
up. About half the total length of when water is available but the
China’s rivers has been lost since Water scarcity infrastructure does not exist to
1990. In North America, the Great There are two types of water utilize it. This is the situation in
Lakes are shrinking, Lake Winnipeg scarcity. Physical water scarcity much of sub-Saharan Africa and
is threatened, and the massive affects regions that naturally do parts of Central America. People
Water stress around the world
KEY
None
1–9 months
10–12 months London

Major cities New York Beijing Tokyo
Istanbul
experiencing Osaka
Los Angeles
water scarcity Cairo Shanghai
Delhi
Mexico City
Mumbai
This map illustrates the

average exposure of water users
to water stress—the ratio of
Rio de Janeiro
total withdrawals to total
renewable supply in a given Sao Paulo
area. A higher proportion of
withdrawals means that more
water users are competing for
limited supplies.

worse. For example, a sewage Salisbury Water
treatment plant run on “clean”
energy can provide the wastewater In Adelaide, South Australia,
an innovative water recycling
needed to fertilize biofuel crops,
system in use in the suburb
which in turn can be used to purify of Salisbury has reduced
The world has not really the water—without emitting extraction from the Murray
woken up to the reality of greenhouse gases. River and aquifers by about
what we are going to face in a half. Wastewater from the
terms of the crises as far Drinkable waste water local sewage treatment works
as water is concerned. New technologies can also convert and rainwater from drains
Rajendra Pachauri wastewater directly into drinkable are treated, and then directed
IPCC Chair water—a process that has been into a series of 50 wetlands.
energy-hungry in the past. The These contain reedbeds
Intergovernmental Panel on Climate and other aquatic vegetation
Change (IPCC) stresses that water that further clean the water.
management policies can lead to The recycled, nonsdrinking
higher greenhouse gas emissions. water from the wetlands is
then piped to the inhabitants
However, that is not the case if the
of Salisbury to use for flushing
living in these areas may have to conversion is fueled using solar
toilets, watering gardens,
spend hours each day walking to energy, which is starting to take washing cars, and filling
the nearest supply of water. Many over from oil to power desalination ornamental ponds.
children miss out on an education plants in the Middle East. In parts In addition to providing
because they are collecting water. of the world, there is seasonal heavy a more sustainable source of
rain—for example, in countries water, the system has boosted
Wildlife concerns with a monsoon—but it runs off biodiversity within the newly
The water crisis is bad for humans into polluted rivers and cannot be established wetlands. Among
and can mean extinction for some used. Rainwater catchment and the birds that are currently
animals and reduction in numbers storage schemes would help. resident or visitors are ducks,
for others. Populations of the Other helpful initiatives include spoonbills, herons, pelicans,
Amazon river dolphin, which lives reducing pollution, cutting irrigation cormorants, and migratory
in the Amazon and Orinoco river and industrial wastage, providing waders, along with species
basins in South America, for new technological solutions for of amphibians and fish, and
many aquatic invertebrates.
example, have been much reduced, developing countries, and reaching
partly by the increase of heavy international agreements—after all,
metal pollution from mining but water catchments do not stick
also by the construction of dams, to national boundaries. ■
which restrict the migration of
fish, the dolphins’ food, to their
spawning grounds. Elsewhere,
in China, the world’s largest
amphibian, the Chinese giant
salamander, has also become
critically endangered by dams There is no water-rich
being built for water storage and country in the world that
hydroelectric power. Such is not facing problems.
engineering works change the Maude Barlow
natural flow of rivers, upsetting Salisbury’s recycled water has
the creature’s habitat. environmental benefits including
reduced demand on existing water
A holistic view of ecosystem resources and improved biodiversity
management is crucial to prevent through the newly created wetlands.
the water crisis from getting much
ENVIRONM
AND CONS
ENTALISM
ERVATION
294 INTRODUCTION
Francis Bacon’s work Written in a cabin UNESCO launch their

espouses the idea that in the woods, Henry The first working Man and the Biosphere
man has dominion David Thoreau’s book photovoltaic solar Programme to encourage
over nature—a view Walden presents a cell panel is built by economic development
that is later termed romanticized view inventor Charles that is sustainable
“imperial ecology.” of the natural world. Fritts in the US. and eco-friendly.
C.1620 1854 1883 1971
1789 1864 1966
Gilbert White’s Natural George Perkins Marsh Lynn White argues that
History and Antiquities of warns of the Western—largely Christian –
Selborne records in great destructive impact anthropocentric worldviews
detail the wildlife around that human action is have placed humankind in an
his rural home. having on nature. environmental crisis.
E
arly in the 17th century, out that resource scarcity was potential for producing limitless
English philosopher and generally the result of human energy, but it took a century for
scientist Francis Bacon actions rather than natural causes. solar power to be widely adopted.
wrote of the need to control and “Clean” hydroelectric power was
manage nature. By the end of the Renewable and clean the first sustainable source capable
18th century, in contrast, English Before the Industrial Revolution, of generating electricity on a large
vicar Gilbert White was writing most energy had been renewable— scale—joined in the late 20th
in favor of a peaceful coexistence the energy of human and animal century by modern wind power, and
between people and the natural labor, wind- and watermills, and tidal, wave, and geothermal energy.
world. Yet in his lifetime, powerful sustainable wood. From the mid-
new steam engines unleashed the 18th century there was a dramatic An environmental ethic
ravages of industrialization—the shift to coal. The most efficient fuel In 1937, following the devastating
reaction against which would later for firing furnaces and factories, it “Dust Bowl” caused by intensive
provide a major impetus for the came at a price—choking pollution farming in the US, President
environmental movement. and the then-unknown rise in Franklin D Roosevelt wrote,
Possibly the first systematic atmospheric greenhouse gases. “A nation that destroys its soils
analysis of humanity’s destructive In the 1880s, however, the key destroys itself.” In 1949, American
impact was American diplomat to a new form of renewable energy ecologist and forester Leopold
George Perkins Marsh’s 1864 was provided by American inventor Aldo articulated a recurring theme
book, Man and Nature. Marsh Charles Fritts—a photovoltaic cell, in environmental thought, by
warned, among other things, which could convert solar power advocating a “land ethic,” a
that deforestation could lead to the to electricity. German industrialist responsible relationship between
creation of deserts, and he pointed Werner von Siemens soon saw its people and their local environment.
ENVIRONMENTALISM AND CONSERVATION 295
The Indian NGO

Navdanya (“Nine The Sustainable Gretchen Daily’s
seeds”) is founded, to Biosphere Initiative Ecosystem Services
preserve seed diversity, (SBI) report, published shows how humans
promote fair trade, and in the US, argues for can derive benefits
protect farmers. increased funding for from preserving the
ecological research. natural environment.
1987 1991 1997
1981 1988 1992 2015
Mark Schafer pioneers The Intergovernmental The UN’s Rio Earth The Paris
Population Viability Panel on Climate Summit sets global Agreement on
Analysis (PVA) as a method Change is established targets for cutting Climate Change
for estimating the likelihood in Geneva, Switzerland. greenhouse gas is signed by 195
of a species’ extinction. emissions. UN countries.
The post-war period saw many 1969 massive oil spill in Santa assets which, when properly
governments legislating to ensure Barbara, California, US senator managed, provide a flow of vital
the quality of air and drinking water Gaylord Nelson proposed the idea goods and services.
and establish national parks and of a national event to highlight the
other protected areas. In 1968, the varied threats to the environment. International cooperation
world first found its collective voice, On the first Earth Day, which took Two UN agencies—the World
when UNESCO (the United Nations place on April 22, 1970, millions Meteorological Organization and
Educational, Scientific and Cultural turned out on marches across the the UN Environment Programme—
Organization) held the Paris US. The scale of the event helped established the Intergovernmental
Biosphere Conference. This resulted, the passage of the Clean Air, Clean Panel on Climate Change (IPCC) in
three years later, in the creation of Water, and Endangered Species 1988 to assess the risk of human-
the Man and Biosphere Programme. Acts and led to the creation of the induced climate change.
United States Environmental The IPCC continues to monitor
Growing awareness Protection Agency (EPA). climate change. In 1992, the Rio
Public concern for the environment In 1973, German economist Earth Summit, a UN initiative, was
was marked by the establishment Ernst Schumacher used the term unprecedented in both its size and
of major conservation organizations. “natural capital” in his best-seller the scope of its concerns. It was the
The International Union for the Small is Beautiful to describe how first of a number of international
Conservation of Nature had been ecosystems provide us with gatherings seeking, with much
established in 1948, and it was complex services. The concept success, to get global agreement on
followed by the World Wildlife Fund inspired American environmentalist greenhouse emissions. International
(1961), Friends of the Earth (1969), Gretchen Daily and others, who cooperation is now seen as key
and Greenpeace (1971). After the argued that ecosystems are capital to saving Earth’s environment. ■
296
THE DOMINION OF
MAN OVER NATURE
RESTS ONLY ON
KNOWLEDGE
HUMANKIND’S DOMINANCE OVER NATURE
T
he Renaissance (“rebirth”) Many scientists of the era believed
IN CONTEXT between the 14th and 17th that humans had a privileged place
centuries is primarily in a universe created by God for
KEY FIGURE
associated with the arts and humanity to inhabit. English
Francis Bacon (1561–1626)
culture that flourished across philosopher and scientist Francis
BEFORE Europe as the Catholic Church’s Bacon (1561–1626), a pioneer in the
c.9500 bce The first authority began to be challenged. It development of scientific method,
agricultural crops are was also a time of extraordinary reinforced this idea; the natural
cultivated in the Middle East. scientific advances, which some world, in his view, existed to
saw as the beginnings of a provide for humans, and should be
340s bce The Greek “scientific revolution.” Discoveries conquered and exploited.
philosopher Aristotle devises a in astronomy, physics, and medicine Bacon’s view later became
“ladder of being” with man at gave rise to the idea that science known as “Imperial Ecology”—the
the top. could tell humans everything about idea that humanity’s knowledge of
the universe, and that knowledge science and technology should be
15th century The “Age of would make humans its masters. used to gain dominance over the
Discovery” begins: Europeans natural world. Imperial ecology
set out to explore the world in became the predominant ideology
search of new resources. throughout the Renaissance, the
AFTER Enlightenment—an 18th century
c.1750 New technology such movement dedicated to the pursuit
as the steam engine launches of knowledge—and later the
Industrial Revolution of the 18th
the Industrial Revolution,
and 19th centuries. ■
which begins in Britain.
1866 Gregor Mendel pioneers Sir Francis Bacon sits for a portrait
the science of genetics, in parliamentary robes. Bacon had
breeding 22 varieties of peas. an illustrious political career; knighted
in 1603, he served as Lord Chancellor
1970s The first experiments of England from 1618 to 1621.
in genetic engineering—the
direct manipulation of DNA by See also: Global warming 202–203 ■ A holistic view of the earth 210–211
humans—take place. ■ Pollution 230–235 ■ Environmental ethics 306–307
NATURE IS A GREAT
ECONOMIST
THE PEACEFUL COEXISTENCE
OF HUMANKIND AND NATURE
I
n the late 18th century, rapid
IN CONTEXT advances in science and
technology—particularly
KEY FIGURE
in Britain—led to widespread
Gilbert White (1720–93)
industrialization and urbanization
BEFORE as people sought to control and From reading White’s Selborne
4th century bce Diogenes, a exploit the natural world. There I took much pleasure in
Greek philosopher, advocates were, however, many in Britain watching the habits of birds,
forgoing the comforts of who still lived and worked on the and even made notes.
civilization in favor of a life land. Among the educated rural Charles Darwin
“in accord with nature.” class, some had a fascination for
both science and nature. From
1773 American naturalist this group, a new generation of
William Bartram starts his field naturalists emerged, suggesting
studies of the wildlife of the that humans should learn from their
southeast US, documented in scientific studies to live in harmony
his 1791 book, Travels. with the natural world rather than onward. The book was compiled
attempt to dominate it. from his correspondence about his
AFTER findings with several like-minded
1949 American ecologist Aldo Arcadian ideology naturalists, but it was more than
Leopold publishes A Sand In 1789, rural parson and naturalist simply a collection of data. White’s
County Almanac, exploring the Gilbert White published his Natural engaging and often poetic style
idea of humans’ “land ethic,” or History and Antiquities of Selborne, sent a persuasive message; his
responsibilities toward nature. which became a seminal work in work rejected the “imperial” idea
what was later called “Arcadian of conquering nature, and instead
1969 Friends of the Earth Ecology”. Educated at Oxford and a encouraged a balance between
is founded in the US—initially keen gardener and ornithologist, humans and the natural world—
as an antinuclear group— White closely observed the wildlife like that of the Ancient Greeks’
marking the beginning of the around his Hampshire village, and mythical idyll of Arcadia, for which
modern Green movement. made meticulous notes from 1751 White’s approach was named. ■
See also: Romanticism, conservation, and ecology 298 ■ Environmental ethics

306–307 ■ The Green Movement 308–309 ■ Halting climate change 316–321
298
IN WILDNESS IS
THE PRESERVATION
OF THE WORLD
ROMANTICISM, CONSERVATION, AND ECOLOGY
I
n many ways, Romanticism— affected scientific attitudes to
IN CONTEXT a new cultural movement that nature by inspiring interest in the
emerged towards the end of nascent field of ecology and the
KEY FIGURE
the 18th century—was a reaction environmental movement.
Henry David Thoreau
to the scientific rationalism of the
(1817–62)
Enlightenment. As industrialization The wild world
BEFORE took hold in urban areas, writers, A key figure in the Romanticization
1662 English diarist John artists, and composers began of nature was Henry David Thoreau,
Evelyn’s work Sylva, advocating increasingly to glorify the natural an American writer from Concord,
forest conservation, is world. The now prosperous middle Massachusetts. His book Walden
presented to the Royal Society. classes were particularly inspired (1854) described his time living in
by Romantic portrayals of nature, a cabin in the woods by Walden
1789 Gilbert White publishes and took up leisure pursuits such Pond. Thoreau advocated
his Natural History of Selborne, as hiking and mountaineering. preserving nature not for its own
inspiring a reaction against The Romantic movement even sake, but as a necessary resource
“imperial ecology.” in sustaining human life and a
kind of spiritual enrichment. While
AFTER Thoreau’s “wilderness” was not far
1872 A bill creating the first removed from modern life, his
US national park, Yellowstone, Romantic portrayal of the natural
is signed into law by President world significantly influenced the
Ulysses S. Grant. conservation movement in the
US and helped inspire the National
1892 In San Francisco,
Parks system. ■
Scottish–American
conservationist John Muir
founds The Sierra Club. Thoreau’s simple hut at Walden
Pond appeared on the title page of
1971 The UNESCO “Man this 1875 edition of Walden. Thoreau
and the Biosphere” project claimed he went to the wilderness
is launched. to be free of the obligations of city life.
See also: Global warming 202–203 ■ A holistic view of Earth 210–211

■ Urban sprawl 282–283 ■ The Green Movement 308–309
MAN EVERYWHERE
IS A DISTURBING
AGENT
HUMAN DEVASTATION OF EARTH
T
he widely held view that
IN CONTEXT the natural world existed to
be exploited by humankind
KEY FIGURE
saw a major rebuttal in the form of
George Perkins Marsh
the 19th-century environmental
(1801–82)
movement. Arguments against
BEFORE the “imperial” attitude to nature,
1824 Joseph Fourier, a French which had prevailed since the dawn
physicist, describes the of global exploration in the late 15th
greenhouse effect—later century, began with naturalists such
identified as a contributing as Gilbert White, and were echoed
factor in global warming. in the sentiments of Romanticism.
Such ideas tended to focus on the George Perkins Marsh in an
1830s Scientists posit that the idealization of nature, rather than engraving from 1882. As well as being
Dutch colonization of Mauritius examining the harm done by human an environmentalist, the Vermont
native was also a skilled linguist,
in the 17th century caused the conquests of the natural world. lawyer, congressman, and diplomat.
dodo to become extinct. In contrast to the emotive
Romantic responses to modernism,
AFTER American polymath George Perkins Marsh believed that people must
1962 In the US, Rachel Marsh took a close look at humans’ be made aware of their destructive
Carson’s Silent Spring impact on the environment and impact and find new ways of
describes the harmful effect of suggested changes. Marsh was managing natural resources to
pesticides on the environment. horrified by the destructive effects preserve the natural equilibrium.
of human management of natural An activist as well as writer, he
1971 Greenpeace is founded
resources. In his book Man and helped establish the principle
by American environmentalists. Nature, Or, Physical Geography as of protected areas, and inspired
1988 The Intergovernmental Modified by Human Action (1864), the idea of sustainable resource
Panel on Climate Change he pointed in particular to the mass management that became a core
(IPCC) is set up to assess the deforestation which had virtually element of the 19th-century
“risk of human-induced desertified some areas of the US. environmental movement. ■
climate change.”
See also: Global warming 202–203 ■ A plastic wasteland 284–285
■ Humankind’s dominance over nature 296 ■ Environmental ethics 306–307
SOLAR
ENERGY
IS BOTH WITHOUT LIMIT
AND WITHOUT
RENEWABLE ENERGY
COST
302 RENEWABLE ENERGY
B
y the late 19th century, fears By contrast, fossil fuels—such
IN CONTEXT were already growing in as coal, oil, and gas—have taken
industrial Europe that the thousands of years to form, and
KEY FIGURE
world could not rely on fossil fuels when exhausted, cannot be
Werner von Siemens
forever. When the first working replaced. Natural gas is an
(1816–92) selenium solar cell panel was built abundant fossil fuel, but its
BEFORE in 1883 by American inventor extraction can cause environmental
2nd century bce The first Charles Fritts, the progressive problems, such as earth tremors
water wheels mark a labor- German industrialist Werner von and water contamination. Nuclear
saving turning point in the Siemens immediately recognized its power, although sustainable for a
history of technology. huge potential for renewable energy. long period of time, is not considered
He declared: “the supply of solar renewable because its production
1839 French physicist Edmond energy is both without limit and requires a rare type of uranium ore.
Becquerel creates the first without cost.” Yet, because no one Energy sources such as solar
photovoltaic cell, using light to at the time understood exactly how power, wind, and water are also
produce a weak voltage. selenium created photoelectricity, generally “clean”—unlike fossil fuels,
and Siemens’s calls for more they produce zero or very low
1873 French inventor Augustin experiments went unheeded, solar greenhouse gas emissions. However,
Mouchot warns that fossil fuels cells were not developed until the not all renewables are clean. People
will run out in the future. 1950s. Today, solar power is the have burned wood and animal dung
1879 The first hydroelectric fastest growing source of new for heat and light for hundreds of
energy and predicted to dominate thousands of years. Trees can be
power plant is built at Niagara
future growth in renewables. replanted and animals produce more
Falls in the United States.
dung, so the practice is sustainable,
AFTER Renewables v. fossil fuels but burning such fuels also emits
1951 Construction of the first Human civilizations have drawn carbon dioxide (CO2), which is one
grid-connected nuclear power on renewable energy for millennia—
plant begins at Obninsk in the from burning firewood to
The Ivanpah solar plant in the
USSR. It produced electricity harnessing the wind to propel Mojave Desert, California, generates
from 1954 to 1959. sailing ships. Renewable sources enough concentrated solar power
such as sunlight or tidal power to serve more than 140,000 homes
1954 Bell Laboratories in the are not at all depleted by use. at peak hours of the day.
US develop the first practical
silicon photovoltaic cell.
1956 American geologist
Marion King Hubbert predicts
declining oil production after
the year 2000.
1966 The world’s first tidal
power station starts operating
on the Rance River in France.
2018 The International Energy
Agency predicts that the share
of renewables in meeting
global energy demands will
increase by a fifth to reach
12.4 percent in 2023.

See also: Global warming 202–203 ■ Pollution 230–235 ■ Ozone depletion
260–261 ■ Depletion of natural resources 262–265 ■ Waste disposal 330–331
Artificial
photosynthesis
Since the early 1970s, scientists
have been working to develop
Solar energy is derived Solar radiation can the technology to mimic the
from solar radiation. be used to produce process of photosynthesis and
energy on Earth. create liquid fuels from carbon
dioxide, water, and sunlight.
All three are plentiful, so if the
process can be replicated it
could produce an endless,
relatively inexpensive supply
The supply of this of clean fuel and electricity.
Solar energy is energy will not stop There are two crucial steps:
without limit. as long as the Sun exists. to develop catalysts that use
solar energy to split water into
oxygen and hydrogen, and
to create other catalysts that
convert hydrogen and carbon
reason why, unlike other forms through an evaporative process,
dioxide into an energy-dense
of renewable energy, they are not first adopted by 16th-century Arab
fuel, such as liquid hydrogen,
classed as “alternative” sources. alchemists and used on an ethanol, or methanol. Scientists
Renewable, clean energy will industrial scale in Chile in the late at Harvard University recently
have huge long-term benefits for 19th century. In the developing used catalysts to split water
populations and ecosystems. It world, solar disinfection is bringing into oxygen and hydrogen,
reduces pollution, mitigates against safe drinking water to more than then fed the hydrogen, plus
global climate change, builds two million people; the process carbon dioxide, to bacteria.
sustainability, and increases the involves using solar heat and ultra- The bioengineeered bacteria
energy security of countries. If it violet light to kill pathogens. converted the carbon dioxide
can be provided cheaply enough, and hydrogen into liquid
it will also pull many people out Wind power fuels. The next challenge is
of poverty. In some 30 countries, For more than 2,000 years, people to transfer a successful lab
renewable energy now makes up have built windmills to pump water experiment into something
more than 20 percent of the supply. and grind grain. Today, wind farms commercially viable.
onshore and offshore account for
Solar energy around 9 percent of renewable
The Sun’s power could supply the energy consumption. A wind
world’s energy needs several times turbine’s huge blades turn around
over. The International Energy a rotor attached to a main shaft,
Agency (IEA) believes that—in which spins a generator to produce
the short term—it has the greatest electricity. Wind power is now the
potential of all the renewables. Its leading area of energy growth in
radiation can be converted directly Europe, the US, and Canada.
into electricity via photovoltaic cells Almost 50 percent of Denmark’s
(as with solar panels on buildings) energy comes from the wind, and
or indirectly by using lenses or in Ireland, Portugal, and Spain the
mirrors to create heat, which can figure is 20 percent. Its global
be converted to electricity. This is potential is thought to be around
called concentrated solar power. five times its present level. This solar fuel generator mimics
the way plants turn sunlight and
Solar panels on a roof can heat It is only economic to build wind carbon dioxide in the air into energy
domestic water. Sunlight can be farms where there is regular wind, and oxygen.
employed to desalinate water however, so the potential is not ❯❯
304 RENEWABLE ENERGY
Hot-dry-rock energy oil, aims to overcome this

problem by fracturing rock
Natural rock fractures that bring strata and injecting water into
hot water to the surface from it at a great depth. The water
deep underground have been is heated by contact with the
described as the “low-hanging rock, then returns to the surface … the wind and the sun
fruit” of geothermal energy through production wells. and the earth itself provide
because they are easy to exploit. Depending on the economic fuel that is free, in amounts
However, they are rare in most limits of drill depth, the that are effectively limitless.
parts of the world. The vast technology might be feasible Al Gore
majority of geothermal energy across many parts of the world, American environmentalist and
locked beneath Earth’s surface but there are risks. Like fracking, former US Vice President
is in dry, nonporous rock. EGS can cause small earth
The enhanced geothermal tremors, so it should not be
system (EGS), a similar process conducted near populated areas
to fracking for natural gas and or power stations.
evenly spread around the globe. Paleolithic times. Ancient Romans close to the surface. The potential
Offshore wind is generally stronger made use of it to heat their villas. is much greater, but drilling for
and more regular than onshore. Today it is employed to generate deep resources is very expensive.
Floating turbines can generate electricity in at least 27 different
wind energy far offshore, unlike countries, with the United States, Water power
seabed-anchored wind turbines, the Philippines, and Indonesia the Since water is 800 times denser
which have to be sited in shallow world’s leading producers. than air, even a slow-moving flow
water close to the coastline. Geothermal heat is also utilized can yield considerable amounts of
directly to heat homes and roads energy if harnessed, for instance,
Geothermal energy in Iceland. Technology is now by dams or tidal barrages that drive
The heat in Earth’s interior is being developed that will use
derived both from the original geothermal hot water to operate
formation of the planet and from desalination plants. The only China’s Three Gorges Dam, the
world’s largest hydroelectric dam, was
the radioactive decay of materials drawback of this renewable energy completed in 2012. Critics point to its
within it. People have bathed in hot source is that it is concentrated ecological impact on the Yangtze River’s
pools, where geothermally heated near tectonic plate boundaries, habitat and biodiversity, and the risk for
water reaches the surface, since where Earth’s mantle heat rises local people of flooding and landslides.

turbines connected to generators. World energy supply by source in 2016
China is the biggest producer of
Hydro
hydroelectric power (HEP), with 2.5%
45,000 small installations in addition
to “big-dam” schemes such as the Natural
Three Gorges project, whose 32 gas
giant turbines have the capacity 22.1%
to produce 22,500 megawatts of
Coal Biofuels
electricity. A downside of big HEP 27.1% and
schemes is that reservoirs created Renewables waste
13.7% 9.5%
upstream of the dam can flood
good farmland, forcing people to
relocate and destroying ecosystems. Oil Nuclear
Despite this, the IEA has estimated 31.9% 4.9%
that by 2023, hydropower will be Others
meeting 16 percent of the global Other
0.3%
renewables 1.7%
demand for electricity.
Tidal power is based on the A pie chart illustrating the sources for the total
same principle: moving water turns energy produced and supplied throughout the world
turbines, which drive electricity in 2016, according to data published by the IEA.
“Others” includes nonrenewable wastes and other
generators. The source of energy sources not included elsewhere such as fuel cells.
from a tidal scheme is reliable,
generating power each time the
tide ebbs and flows, but such of Scotland in 2000, and the first grain for biofuels, can also damage
schemes are expensive to construct. multi-generator wave farm opened the environment. Perhaps because
At present, the largest is the Sihwa at Aguçadoura in Portugal in 2008. of this, biomass is a more common
Lake Tidal Power Station in South fuel in nations that cannot afford
Korea, which was completed in Biomass other renewable options. According
2011 and has reduced the annual Organic matter from plants or to the IEA, the majority of solid
amount of CO2 the nation generates animals is known as biomass. biofuel supply in 2016 took place in
by 315,000 tons (286,000 tonnes). It contains stored energy because Africa, accounting for 33.2 percent.
Wave power involves the capture plants absorb the solar power they
of wave energy through a converter. need for growth via photosynthesis, The future
The first commercial wave power and creatures absorb that energy As growth in renewables increases,
scheme began off the west coast either from the plants they eat the advantages of each type must
or from what their prey consumes. be balanced against their adverse
Creating a renewable fuel from effects—from biomass pollution to
plant, animal, and human waste the reported role of wind turbine
products such as straw, dung, and blades in the deaths of migrating
garbage may seem an attractive birds. In 2014, the IEA predicted that
… someday, renewable energy option, and some coal-fired power renewables would provide 40 percent
stations have been converted to of global energy needs by 2040. In
will be the only way for people
wood-burners. Burning biomass 2018, the IEA further predicted that
to satisfy their energy needs. produces heat, electricity, and renewables would account for almost
Hermann Scheer transport fuels, such as ethanol and a third of all world electricity by
President, European Association
for Renewable Energy biodiesel. However, biomass energy 2023, with solar power taking the
is not necessarily “clean.” Burned biggest share. Energy from ocean
as a fuel, biomass releases CO2, and currents could also generate huge
creates air and particle pollution. amounts of electricity, as could
Clearing prime forest for its wood or large arrays of solar panels in space
to cultivate biomass crops, such as or floating on the seas. ■
306
THE TIME HAS COME

FOR SCIENCE TO BUSY
ITSELF WITH THE
EARTH ITSELF
ENVIRONMENTAL ETHICS
A
t its heart, the discipline documented the serious impact
IN CONTEXT of environmental ethics of pesticides on the environment,
extends the boundaries and brought these issues to the
KEY FIGURE
of ethics beyond humans, and into forefront of American public debate.
Aldo Leopold (1887–1948)
the natural world. It forces humans Six years later, US ecologist Garett
BEFORE to question their role in the Hardin’s article “The Tragedy of the
1894 In The Mountains of environment, their responsibility to Commons” outlined the danger of
California, Scottish–American the planet itself, and their duty to overusing shared resources and
naturalist John Muir describes future generations. allowing the human population
his travels through wild places The field of environmental to grow unchecked.
in California, evoking the deep ethics grew out of an urgent sense Other writers viewed the
spirituality and adventure he of encroaching crisis, expressed impending crisis from a more
feels when in the wilderness. in both popular and academic philosophical perspective. Aldo
writings. In 1962, the book Silent Leopold’s “land ethic,” outlined in
1909 Gifford Pinchot’s The Spring, written by US biologist and A Sand County Almanac (1949),
ABC of Conservation argues conservationist Rachel Carson, placed human beings on an equal
that future generations should footing with other species in a
be able to utilize Earth’s wider ecosystem. As one part of a
natural resources. larger whole, our ethical concerns
should be with the healthy
AFTER functioning of the entire ecosystem,
1968 US academic Paul R. rather than simply the advancement
Erlich and his wife, Anne, A thing is right when of human health and happiness.
publish The Population Bomb, it tends to preserve the In his seminal 1966 lecture
warning of the dangers of integrity, stability, and “The Historical Roots of Our
human population growth. beauty of the biotic Ecologic Crisis,” later published
community. It is wrong as an article, the US historian
1970 On April 22, the first when it tends otherwise. Lynn White claimed that the
Earth Day is celebrated in the Aldo Leopold environmental crisis was the fault
US. It becomes an annual of Western society’s worldview.
global celebration of In particular, he blamed the
environmental education Christian thinking that promoted
and reform. anthropocentrism—the idea that
humans are superior to all other

See also: Endangered habitats 236–239 ■ Pesticides 236–239 ■ Depletion of
natural resources 262–265 ■ Ecosystem services 328–329
Aldo Leopold
Born in 1887, Aldo Leopold
grew up in Burlington, Iowa.
He received his degree from
the Yale School of Forestry,
after which he took a job with
the US Forest Service. While
there he was instrumental
in the proposal to manage
The remote, subalpine Mineral wanted to build a ski resort there. the Gila National Forest as a
King Valley has survived the threat of The Valley had no official protected wilderness area, and in 1924
development. It remains an ecosystem it became the first official
designation beyond that of a game
that aims to benefit all—following Aldo Wilderness Area in the US.
refuge, but the Sierra Club argued
Leopold’s “land ethic” principle. Leopold then moved to
that the area should be preserved Wisconsin to continue his work
in its original state for its own in the Forest Service, and in
creatures, leading to the view that sake. The suit went to the Supreme 1933 became a Professor
nature was created for humanity’s Court, which in 1974 ruled in favor of Game Management at the
use and exploitation. of the Forest Service and Disney. University of Wisconsin.
By then, however, Disney’s interest Leopold died in 1948 while
Ethical dilemmas had waned; today the Valley is part helping fight a grass fire.
Environmental ethics questions of Sequoia National Park. Most of his many essays
the moral imperatives behind The battle between those who on natural history and
sustainability and stewardship by follow anthropocentric ethics and conservation were published
asking if the motivations are those who argue for ecocentric posthumously in collections,
grounded in anthropocentrism, or approaches has continued. It often such as A Sand County
in the protection of the natural world takes place in political arenas, Almanac, that greatly
influenced the emerging
because it inherently deserves particularly with the increased
environmental movement.
protection. These questions have prominence of globally sensitive
played out not only in philosophical issues such as climate change.
Key works
arenas, but also in the legal and Sustainable development has
political spheres. generally been an anthropocentric 1933 Game Management
In 1969, the Sierra Club, an endeavor, to ensure future 1949 A Sand County Almanac
environmental lobbying group, generations have their needs met. 1953 Round River: From the
challenged a US Forest Service Environmental ethicists tend to Journals of Aldo Leopold
permit allowing Walt Disney argue that sustainability is only 1991 The River of the Mother
Enterprises to survey the Mineral viable if it preserves the future of all of God: and Other Essays
King Valley in California—Disney members of the ecosystem. ■
308
THINK GLOBALLY,
ACT LOCALLY
THE GREEN MOVEMENT
T
he roots of the modern its height and the Cuban Missile
IN CONTEXT “green movement” Crisis of 1962 brought the US and
developed in organizations the Soviet Union to the brink of
KEY FIGURES
established in the late 19th and nuclear war, galvanizing calls for
David Brower (1912–2000),
early 20th centuries, such as the nuclear disarmament among
Petra Kelly (1947–92)
Sierra Club. Faced with the threat many campaigners.
BEFORE of increasing urbanization and In this atmosphere, the idea
1892 The Sierra Club is industrialization, the Sierra Club of conserving particular natural
founded in San Francisco, sought to protect the natural sites, as in the national parks
California, by the Scottish– environment for people’s enjoyment. system in the US and the UK,
American conservationist A greater awareness of humans’ gave way to a much broader
John Muir. relationship with the environment concept of environmentalism.
led to the emergence of a more Several organizations emerged with
1958 Environmentalists politically active environmental a strong activist agenda involving
protest against proposals movement in the second half of the mass protests and direct action.
for a nuclear power plant 20th century. This took off in the
at Bodega Bay, California. 1960s, when the Cold War was at Organized protest
One of the first of the activist
AFTER organizations was Friends of
1970 On April 22, the first the Earth. It was founded in the US
Earth Day is held across the US. in 1969 by a group that included
1972 Environmentalist conservationist David Brower,
a former leader of the Sierra Club,
candidates stand for election Only through care with the aim of preventing the
in Tasmania, New Zealand, for the environment can building of nuclear power plants.
and Switzerland. the livelihoods of those Politically active from the outset,
1996 Ralph Nader stands as most dependent on it Friends of the Earth continues to
candidate for President of the be sustained. lobby governments across the world
US on the Green Party ticket. Petra Kelly and campaigns on a broad range of
environmental issues, emphasizing
the importance of sustainable
economic development. In 1971,
a small group of activists in North
America formed the Don’t Make a

See also: Citizen science 178–183 ■ Pesticides 242–247 ■ Human devastation
of Earth 299 ■ Halting climate change 316–321
Petra Kelly
Born Petra Lehmann in
Günzburg, West Germany,
in 1947, Kelly later adopted
the surname of her stepfather,
an American army officer.
When she was 12, the family
moved to the US, where Kelly
studied political science in
Activists in a dinghy patrol in front formed in Germany in 1979. As Washington, D.C.
of two ships from the UK carrying the movement gained momentum, In 1970, Kelly returned
illegal toxic substances as part of to Europe. While working
many smaller parties began to
regular Greenpeace protests. at the European Commission
coalesce to form national, unified
in Brussels, she joined
Green Parties. Germany’s Social Democratic
Wave Committee to protest against In recent years, as issues such Party, but grew disillusioned
nuclear bomb testing by the US on as pollution and climate change with traditional politics. She
the island of Amchitka, Alaska. have risen up the news agenda, joined Germany’s newly
The organization favored direct other established political parties formed Green Party in 1979,
action rather than political lobbying have adopted environmentally and in 1983 was one of 28
and chartered a boat to sail to the friendly policies. ■ members to be elected to
island in protest. The publicity parliament. Kelly campaigned
generated by the group swayed on issues of environmentalism
public opinion and halted the tests. and human rights. In 1992,
This was the first action of what she and her companion, Green
was to become Greenpeace, an politician Gert Bastian, were
organization that continues to found dead at her home in
We have everything Bonn, apparently the result
use direct action to challenge
we need, save perhaps, of a suicide pact.
those engaged in environmentally
damaging activities. political will. But, you know
what … political will is a Key works
Green politics renewable resource.
Al Gore 1984 Fighting for Hope
During the 1970s, political parties
1992 Nonviolence Speaks
with dedicated environmentalist
to Power
manifestos emerged in several 1994 Thinking Green: Essays
countries. For example, The Ecology on Environmentalism,
Party was established in the UK Feminism, and Nonviolence
in 1975, and the Green Party
310
THE CONSEQUENCES OF
TODAY’S ACTIONS ON
TOMORROW’S WORLD
MAN AND THE BIOSPHERE PROGRAMME
D
uring the second half of UNESCO was founded after World
IN CONTEXT the 20th century, there was War II with the aim of fostering
an increasing global “the building of peace, the
KEY ORGANIZATION
awareness of the importance of the eradication of poverty, sustainable
UNESCO
relationship between humans and development and intercultural
BEFORE the natural world. This led, in dialogue through education, the
1925 The International 1971, to the United Nations sciences, culture, communication
Institute of Intellectual Educational, Scientific, and Cultural and information.” As such, it was
Cooperation—which aims to Organization (UNESCO) launching in a unique position to examine
exchange intellectual ideas the Man and the Biosphere carefully the relationship between
and improve quality of life—is Program (MAB). This is an people and the environment.
set up in Paris, France. intergovernmental program devoted
to encouraging environmentally Global network
1945 The United Nations sustainable and equitable economic The organization began by setting
Conference establishes the development, while protecting up a number of internationally
constitution of UNESCO. natural ecosystems. recognized protected sites, known
AFTER
1983 First International
Biosphere Reserve Congress
takes place in Minsk, Belarus. Humankind is altering the
environment with processes Such actions have
1995 Statutory framework such as deforestation and consequences.
of the World Network of urban sprawl.
Biosphere Reserves is agreed.
2015 The UN launches its
17 Sustainable Goals initiative.
2017 The US withdraws The MAB program
17 sites from the UNESCO predicts the Data gathered from global
MAB reserves helps
World Network of Biosphere consequences of generate a picture of what
Reserves, but 23 new sites today’s actions on these consequences could be.
are added elsewhere. tomorrow’s world.

See also: Human activity and biodiversity 92–95 ■ The ecosystem 134–137 ■ The peaceful coexistence of man and
nature 297 ■ Renewable energy 300–305 ■ Environmental ethics 306–307 ■ Sustainable Biosphere Initiative 322–323
as the World Network of Biosphere

Reserves (WNBR). These set out
to show how human cultural and
biological diversity are mutually
beneficial and encourage the
balanced integration of people with
their natural environment. They
also sought to find ways to manage
natural resources efficiently for the
benefit of the environment as well
as its inhabitants.
There are now over 650 sites
around the world, providing a
platform for collaborative scientific
and cultural research in a range
of marine, coastal, and terrestrial
ecosystems. Through the network,
the program monitors the effects of achieved by zoning areas within Moroccan women gather the
human activity on the biosphere, the reserve to protect core health-giving fruits of the argan
particularly examining climate locations, whilst simultaneously tree. These trees in the Arganeraie
Biosphere Reserve are carefully
change, and fosters the exchange providing places for appropriate sustained by the local population.
of information. and sustainable development by
local inhabitants.
Local knowledge To this end, communities are sharing knowledge across the
The MAB program recognizes encouraged to participate in the World Network is key to the
three interconnected functions of management of the reserve, and success of the project as a whole.
a biosphere reserve: conservation; use their local knowledge of the
sustainable development; and area to make the best use of natural Conflicting opinions
support though education and resources. The idea of educating The sites of the WBNR, as well as
training. These objectives are people about the environment and being of international scientific
significance, are often culturally
UNESCO the outbreak of World War II. important to the host state. They
Today, members aim to achieve are not nominated by UNESCO, but
UNESCO, an agency of the UN their objectives by sponsoring by national governments, and they
based in Paris, France, was international educational and remain under the jurisdiction of the
founded in 1946 to promote scientific programs. These states they are in. International
international collaboration for include dedicated projects recognition of their status does not
peace and security. It was that promote and protect impinge upon the rights of those
established in line with the human rights and sustainable states over the Biosphere Reserves.
United Nations Charter, through development, while encouraging In recent years, some states
education, science, and culture. cultural diversity. have chosen to manage certain
Today, the organization has 195 The organization is perhaps sites as national rather than
member states. best known for establishing
international reserves and have
UNESCO continues the work internationally recognized World
withdrawn them from the program.
begun by the League of Nations Heritage Sites, which aim to
International Committee on preserve as many aspects as Nevertheless, there has been a
Intellectual Cooperation in the possible of the world’s diverse steady increase in sites nominated
1920s, which was interrupted by cultural and natural heritage. for the program from governments
around the world. ■
312
IN CONTEXT
PREDICTING
KEY FIGURE
Mark L. Shaffer (1949–)
A POPULATION’S
BEFORE
1964 The IUCN publishes its
first Red List of threatened
mammal and bird species.
SIZE AND ITS

1965 In The Destruction of
California, ecologist Raymond
Dasmann charts the rapid loss
CHANCES OF
of flora and fauna in the state.
1967 The Theory of Island
Biogeography by Robert
MacArthur and Edward O.
EXTINCTION
Wilson explores island patterns
of immigration and extinction.
AFTER
2003 Population viability
POPULATION VIABILITY ANALYSIS analysis (PVA) of the Fender’s
blue butterfly is used to guide
conservation in the US.
2014 PVA studies in the
Sonoran Desert, US, help
assess the response of birds
and reptiles to climate change.
P
opulation viability analysis
(PVA), or extinction risk
assessment, is a process
used to estimate the probability
that a population of a target species
has the ability to sustain itself for
a specific time, be it 10, 30, or 100
years. A key feature of PVA is the
definition of minimum viable
population sizes and minimum
habitat areas—information which
can then inform decisions on
conservation priorities.
A tool for conservationists

PVA combines both statistics and
ecology to calculate the fewest
organisms required for a species
to survive long-term in its preferred

See also: Ecological resilience 150–151 ■ Climax community 172–173
■ Metapopulations 186–187 ■ Mass extinctions 218–223 ■ Deforestation 254–259
Vulnerability of
small populations
Fender’s blue butterfly was not seen A minimum viable population
after the 1930s and was deemed extinct has to be of a sufficient size
until it was rediscovered in 1989. It is not only to maintain itself
endangered, but small populations live under average conditions but
in northwest Oregon. also to endure extreme events.
Mark Shaffer likened this to
a reservoir built to withstand
habitat, it takes no more than a low the type of flood that occurs
level of environmental damage or only once in 50 years, but not
human disturbance to nudge them a devastating once-in-a-
toward extinction. century flood.
Small populations are
Counting grizzlies especially vulnerable to
In 1975, grizzly bear numbers were multiple threats occurring
shrinking in Yellowstone National successively. The Heath Hen
Park. Only an estimated 136 of the in New England, US, had been
bears were left, and this isolated widespread in colonial times,
but relentless hunting for food
habitat. This minimum number population was considered to be
and sport caused a dramatic
also dictates the amount of suitable endangered. As part of his doctoral decline in Heath Hen numbers
habitat that the species needs. PVA research, Mark L. Shaffer began to by 1908. In that year, the last
is a useful tool for conservationists study the long-term sustainability surviving population on the
when lobbying governments and of this geographically isolated island of Martha’s Vineyard
developers to give protected status grizzly bear population. was given protected status.
to an area. Armed with a PVA, they Shaffer, a pioneer of population However, a catastrophic
can explain precisely why reducing viability analysis, applied four wildfire during the 1916
a stretch of forest, heathland, or factors that he considered would breeding season, severe
reedbed will threaten certain flora decide their fate. The first was winters, inbreeding, disease,
or fauna. Protecting an area that demographic stochasticity: and heavy predation by birds
is extensive enough to support irregular, unpredictable fluctuations of prey all combined to push
a large species also benefits many in numbers, age, gender, and birth the Heath Hen population
smaller organisms sharing the and death rates. For example, if the below a viable level. By 1927,
same environment. overwhelming majority of animals only two females remained,
and the species was extinct
A number of creatures can only in a population are males, breeding
by 1932.
survive in environments where success will be poorer than in a ❯❯
human disturbance is minimal.
This is especially true for those
that live in specialist habitats,
such as certain owls in old-growth
forest, reptiles on acid heathland,
or amphibians in fast-flowing,
unpolluted streams. However, as Uncertainty is just about
the human population grows, there the only certainty in PVA.
is a constant demand for land for Steven Beissinger
building, agriculture, leisure, roads, American conservation biologist
or forestry. This pressure is a
particular threat to species that
cannot easily adapt and move
elsewhere. Where they are already
confined to “islands” of suitable
314 POPULATION VIABILITY ANALYSIS
A female grizzly and her cubs forage extensive Greater Yellowstone modllers have suggested that
in Yellowstone. A female’s home range Ecosystem—an area of 34,375 sq Yellowstone may have reached its
is 300–550 sq miles (775–1,400 sq km), miles (89,031 sq km) that has the maximum carrying capacity—the
while a male’s is as much as 2,000 sq
miles (5,000 sq km).
national park at its core. In 2014, largest number of animals an area
the US Geological Survey estimated of suitable habitat can support. In
that around 757 bears lived in the 2017, grizzlies were briefly removed
more evenly balanced population, ecosystem, based on 119 sightings from the threatened species list,
and will influence its chances of of grizzly sows and cubs. However, but their protections were restored
survival. The second consideration the population had dropped to by a federal judge in 2018.
was environmental stochasticity: around 718 in 2018, and population
unpredictable fluctuations in How studies are devised
environmental conditions, such PVA studies are now conducted in
as habitat and climate changes, several ways. The simplest type is
which may affect the availability the time-series PVA, which looks at
of food and shelter. The third the entirety of a population over a
was natural catastrophes, such period of time in order to calculate
as forest fires or floods. The fourth Technology is increasingly a rough average growth trend and
of Shaffer’s factors was genetic allowing scientists and any variations. In such studies, all
changes, including problems policymakers to more closely individuals are treated as identical.
created by inbreeding. For each monitor the planet’s Demographic PVAs tend to be
of these, statistical modeling can biodiversity and threats to it. more precise and detailed. They are
determine a range of possibilities. Stuart L. Pimm based on estimated reproductive
Since Shaffer’s initial research American–British biologist and survival rates for different age
in the 1970s and ’80s, and bands within the population. Such
subsequent new management and analyses require much more data,
conservation strategies, grizzlies but can provide extra information
have extended their habitat by on the needs and vulnerability of
more than 50 percent within the different sections of the population,
A population is identified as being at risk.
A population A management
viability analysis is solution is found to
conducted to assess combat the threat
the situation. to the population.
The island foxes of the Channel

The population has Islands, off California, numbered fewer
than 200 in the late 1990s. By 2015,
a chance to recover. there were more than 5,000, but on one
island, a subpopulation is still at risk.
fueling a case for conservation “surrogate” data from one colony dolphins off the coasts of Argentina
where protection is required. As was used to make forecasts on the and Australia. With the development
reliable information on age ranges other two; they proved valid for one of increasingly efficient computer
and breeding rates is often not colony, but not the other. programs incorporating ever more
available for small, threatened variables, PVA will undoubtedly
populations, ecologists sometimes Making a difference be used even more effectively in
use data from other populations of Methods are still being refined, but the future. It is impossible to predict
the same species—or a different but PVA has now become a cornerstone every extinction, but PVA provides
similar species—to conduct a PVA. of conservation biology. PVAs have tools for identifying endangered
However, the results are variable, been applied to populations as populations and determining the
even in populations of the same varied as island foxes in California, management actions likely to
species in the same area. In a 2015 sea otters in Alaska, Fender’s blue be most effective in improving
study of three colonies of California butterflies and Northern Spotted population viability, and preserving
sea lion in the Gulf of California, Owls in Oregon, and bottlenose a species at risk. ■
A Japanese study collected population growth

data, including the number of
The Japanese Rock Ptarmigan female offspring that survived
lives in the Japanese Alps at to the next breeding season and
an altitude of around 8,200 ft the annual survival rate of all
(2,500 m). Its population of some birds. Their calculations included Population viability
2,000 birds is divided into variables for a range of offspring analysis can indicate how
several small communities from each pair. urgently recovery efforts need
on mountain peaks. When a Their findings indicated that to be initiated in specific
combination of climate warming there was a relatively low risk of populations.
and predators moving further extinction in the next 30 years, William F. Morris
up the mountains prompted even if the starting population American biologist
fears for its survival, ecologist was only 15. One potential
Ayaka Suzuki and his team set conclusion is that the Mount
out to find the minimum viable Norikura population is strong
population size for the birds on enough to supplement declining
Mount Norikura. The team populations on other mountains.
CLIMATE
CHANGE
IS HAPPENING HERE.
IT IS HAPPENING
HALTING CLIMATE CHANGE
NOW
318 HALTING CLIMATE CHANGE
S
ince the Industrial
IN CONTEXT Revolution, humans have
been altering Earth’s natural
KEY FIGURES
environment through increased
Bert Bolin (1925–2007),
carbon dioxide (CO2) emissions.
Intergovernmental Panel Societies have become more
on Climate Change (1988–) technologically advanced, but this
BEFORE technology—from coal-powered
1955 American scientist trains, ships, and factories, to oil-
Gilbert Plass concludes that fueled cars and planes—has had
higher concentrations of an adverse impact on the natural
carbon dioxide (CO2) will lead world and the species inhabiting
to higher temperatures. it. As scientists have become more
aware of the human causes of
1957 American scientist Roger climate change, global research
Revelle and Austrian physical groups have been formed to study
chemist Hans Suess jointly the phenomenon and suggest ways
publish a report proving that in which humanity can halt, if not Firefighters battle flames from the
the oceans will not absorb the reverse, the damage. “Holy Fire” that ravaged Orange
The effects of climate change County, California, in 2018. Higher
excess CO2 in the atmosphere. temperatures led to an extended and
are varied. As more CO2 in the
1968 British glaciologist difficult forest fire season.
atmosphere creates global warming,
John H. Mercer theorizes a this causes the polar ice caps to
catastrophic rise in sea levels melt, the oceans to warm and rise, related catastrophes, such as those
in the next 40 years due to the and species that are unsuited to impacted by tropical monsoons,
collapse of Antarctic ice sheets. warmer oceans to die out. Global are seeing the most severe
weather patterns are also changing: repercussions, especially
AFTER hurricanes in the North Atlantic in terms of loss of life and habitat.
2020 Plans created by the region have increased in intensity,
Paris Agreement to combat leaving devastation and death in Global cooperation
climate change are due to their wake. Fires and droughts have Scientists have been aware
be implemented. become more frequent in dry areas; that human actions contribute
winters are more severe in colder to climate change since 1896, when
climates. Areas of the world already Swedish scientist Svante Arrhenius
susceptible to extreme weather- suggested that people burning
Natural causes: Human causes:

• Volcanic eruptions • Deforestation
• Shifting plate tectonics • Farming practices
• Ocean currents • Fossil fuel burning
• Industrial emissions
... lead to increased amounts of

carbon dioxide in the atmosphere,
causing climate change.

See also: Global warming 202–203 ■ Deforestation 254–259 ■ Man and the Biosphere Programme 310–311
■ Sustainable Biosphere Initiative 322–323 ■ The economic impact of climate change 324–325
fossil fuels were adding to global agreement, which was ratified by

warming. It was not until the 1970s, all UN member states, did reduce
however, that governments began greenhouse gas emissions.
to act upon this knowledge. Around
this time, the general public had Creation of the IPCC
begun to be made aware of the … human beings are now In 1988, the Intergovernmental
reality of climate change due to carrying out a large-scale Panel on Climate Change (IPCC)
news articles and broadcasts that geophysical experiment of a was established in Geneva,
shared the bleak outlooks of climate kind that could not have Switzerland, by two United Nations
scientists with the wider world. happened in the past… organizations: UNEP, and the
International efforts to halt or Roger Revelle and World Meteorological Organization
delay climate change began with Hans Suess (WMO). Swedish meteorologist
the first United Nations conference Bert Bolin—who served on the
on the environment, which was Advisory Group on Greenhouse
held in Stockholm, Sweden, in Gases that the IPCC supplanted—
1972. The conference paid little was the panel’s first chairman.
attention to the issue of climate The IPCC was created to serve
change compared to other as a globally coordinated response
environmental issues—such as ecosystem management, natural to climate change linked to human
pollution and renewable energy— disaster relief, and antipollution activity. It issues reports based on
but did create the United Nations activities. UNEP later became scientific research in support of
Environment Programme (UNEP), responsible for coordinating UN the main international treaty on
an agency to oversee environmental efforts against climate change. climate change: the UN Framework
policies and programs such as In 1987, UN members also Convention on Climate Change
agreed to the Montreal Protocol, (UNFCCC), which was signed at
pledging to protect Earth’s ozone the Earth Summit in Rio de Janeiro,
An Inconvenient Truth, a 2006
documentary on climate change layer by ending the use of ozone- Brazil, in 1992. The IPCC’s work
by former US Vice President Al Gore, depleting substances. Although also involves issuing the Summary
aimed to educate the public on the it was not specifically designed for Policymakers (SPM), which
causes and effects of climate change. to combat climate change, the provides summaries of climate ❯❯
320 HALTING CLIMATE CHANGE

change research to governments
around the world to help them
understand the threats to humans
and the environment as a result of
climate change.
The Kyoto plan

Nine years after the creation of the
IPCC, in 1997, UN members signed
the Kyoto Protocol, which sought to
improve regulation of global carbon
emissions. This protocol was the
first agreement among nations
to mandate country-by-country
reductions in greenhouse gas
emissions, aiming to reduce them
to levels that would stop humans
from negatively impacting the
world’s ecosystems.
Although signed in 1997, the are on track to meet their target for An underwater cabinet meeting is
Kyoto Protocol did not take effect 2020, except for Norway, which had held in the Maldives in 2009 to call for
until 2005. At the end of the first set a very high target (a 30–40 action against climate change. Rising
sea levels could mean that the nation
commitment period in 2012, all percent reduction from 1990 levels). is eventually swallowed by the ocean.
signatory nations had achieved
their target reduction except for Paris and the future
Canada, which withdrew from the The Kyoto Protocols set targets for more aggressive global resolution of
protocol because it could not meet nations to meet from 2005 to 2020. climate change, the Agreement was
its targets. Australia also failed to After 2020, signatory nations will signed by 195 UNFCCC member
reduce emissions, but in the initial begin to abide by a new protocol: countries at the UN headquarters
period, their target was set as an the Paris Agreement. In November in New York City. Like Kyoto, the
8 percent increase. Most nations 2016, after decades of calls for a primary aim of the Paris Agreement
is to cut greenhouse gas emissions
Climate change denial to agreed-upon levels.
With the decision of Syria to
Despite the consensus by the Some scepticism comes from sign the Paris Agreement in 2017,
majority of scientists around the those who suggest scientists’ the United States became the only
world that climate change is a estimations are too alarmist, country in the world not to take
human-caused phenomenon and and that global warming is part in the agreement. Although the
requires urgent intervention, happening more slowly than US initially signed the agreement
climate change denial persists predicted. Others see the idea under Barack Obama’s presidency,
in many of the world’s most of climate change as a human his successor, Donald Trump, has
powerful nations. Several phenomenon as a hoax, instead rejected the agreement, claiming
scholars have termed the claiming that global warming is that it asked too much of the United
opposition to the facts of climate a natural cycle for the planet States and too little of other nations.
change a “denial machine,” in and not a product of human
This decision struck a blow to the
which conservative media and behavior. Whatever the reason,
industries benefitting from lax denial of climate change among other signatories; as well as having
environmental regulations some policymakers and business plenty of wealth to fund climate
create an environment of leaders is a position that that research, the US is also the world’s
uncertainty and scepticism the IPCC and scientists continue second-largest greenhouse gas
about climate change science. to disprove. emitter. President Trump has since
clarified his position by saying

that he believes climate change The burden of change
to be a natural phenomenon from
which the world can “come back”
without significant changes to
human behavior.
Other nations have voiced their Developed nations
Greenhouse gas
own concerns with the Paris emissions must
will take the
lead in reducing
Agreement. The government of peak as soon
carbon emissions.
Nicaragua, which joined the accord as possible.
in 2017, criticized the Agreement
for not going far enough and argued
that it will not reduce carbon
emissions quickly enough to avert
The losses suffered
global climate disaster. The Paris by vulnerable Developed
Agreement also lacks a mechanism nations due to countries will
climate change provide financial
to ensure that countries that have help to developing
must be addressed.
signed it comply with its terms. countries.
Desperate measures
According to the terms of the Paris
Agreement, countries must work
together to limit the increase in The Paris Agreement was signed by 195
the global average temperature member countries of the UNFCCC. It placed
to below 2°C (3.6°F) above pre- the responsibility on developed nations to assist
those who lacked the funds or resources to
industrial levels. The Agreement combat climate change alone.
also seeks to go further, suggesting
that the increase should be limited
to only 1.5°C (2.7°F). In a study create a global environment create no emissions without also
published in the journal Earth mirroring the current highest removing an equivalent amount of
System Dynamics in 2016, climate temperatures experienced, a 2°C CO2 from the atmosphere.
scientist Carl-Friedrich Schleussner increase would usher in a “new The IPCC’s 2018 report also
and his co-researchers argued that climate regime” unlike anything appealed to individuals to do their
while an increase of 1.5°C would humans have seen before. part to lower CO2 emissions. Land
Subsequent research has shown use, energy, cities, and industry are
that this 1.5°C target will prove the major areas in which the IPCC
difficult to meet. In 2018, the IPCC suggests change is necessary:
produced a Special Report on global people should embrace electric cars;
warming, as it had been tasked to walk and bicycle more; and fly less
do by the Paris Agreement. Its often, because planes produce a
We have presented findings were alarming. Rather significant proportion of greenhouse
governments with pretty than being on track for the 1.5°C gases. The IPCC also encouraged
hard choices. We have pointed target, the world is now headed people to buy less meat, milk,
out the enormous benefits of closer to 3°C above preindustrial cheese, and butter, because reduced
keeping to 1.5°C. levels. To recover and hit the target demand for these products should
Professor Jim Skea of 1.5°C would require nations to lead to lower emissions by the meat
Co-Chair, IPCC working group III take unprecedented and drastic and dairy processing industries.
measures. Global human CO2 While global agreements such as
emissions would need to drop 45 Kyoto and Paris have dominated the
percent from 2010 levels by 2030, conversation, it is now clear that
and in 2050, would need to reach any and all methods to lower CO2
“net zero,” meaning that humans emissions must be pursued. ■
322
THE CAPACITY
TO SUSTAIN THE
WORLD’S POPULATION
SUSTAINABLE BIOSPHERE INITIATIVE
T
he Sustainable Biosphere useful ecological knowledge as
IN CONTEXT Initiative (SBI) emerged in scientists raced to combat
1988 due to the efforts of environmental degradation.
KEY FIGURE
the Ecological Society of America
Jane Lubchenco (1947–)
(ESA) to establish what scientific Prioritizing the planet
BEFORE research should be prioritized given The scientists of the SBI set out a
1388 England’s Parliament the limited funding available. new path for the field of ecology,
makes it illegal to throw waste At this time, the field of ecology and determined which research
into public watercourses such was undergoing a transition areas would be the most important
as ditches and rivers. towards applied science—using in the years to come. They sought
knowledge to develop practical to prioritize three fields of research:
1970s British scientist James solutions relevant to contemporary global change, biological diversity,
Lovelock and American environmental issues. American and sustainable ecological systems.
microbiologist Lynn Margulis environmentalist Jane Lubchenco Studies of global change look at the
develop the Gaia hypothesis. led the SBI, and paved the way atmosphere, climate, soil and water
for the ESA (and others) to promote (including changes due to pollution),
AFTER and patterns of land- and water-use.
1992 Canadian ecologist Research into biological diversity
William Rees introduces the focuses on the conservation of
concept of the “ecological endangered species and the study
footprint” to describe human of natural and manmade changes in
impact on the environment. genetic and habitat diversity. Finally,
The SBI has stimulated studies of sustainable ecological
2000 Dutch Nobel laureate improvements in systems analyze the interactions
Paul Crutzen popularizes the understanding and in between humans and ecological
idea that the world has entered advancing connections processes in order for scientists to
a new geological epoch known between ecological find solutions to the stresses they
as the Anthropocene, or “Age knowledge and society. detect in ecosystems.
of Man.” This era recognizes Jane Lubchenco The SBI stressed the need for
the monumental and often funding for such research, and also
dangerous ecological impacts highlighted the importance of
humans make on the planet. sharing findings with those outside
the scientific community. It set out
a process for applied ecological

See also: The ecosystem 134–137 ■ Chaotic population change 184 ■ Gaia 214–217 ■ Overfishing 266–269
■Halting climate change 316–321 ■ The economics of climate change 324–325 ■ Waste disposal 330–331
research that included not only

acquiring new knowledge, but
communicating it and helping
incorporate it into real-world policy
changes.
The future of research

Lubchenco and her colleagues
created the SBI as both a mission
statement and an argument for why
ecological research deserved more
funding and attention. Their report
was published in 1991 in the journal
Ecology as “The Sustainable
Biosphere Initiative: An Ecological
Research Agenda.” It was well
received within the scientific
community, and has been adapted workshops, and creating reports to Wind turbines are explained to young
for use at a global level—first in the advance its agenda. The SBI has students. The SBI advocates ecological
International Sustainable Biosphere brought ecology into the public eye, education in schools and universities so
that people can learn how to manage
Initiative that was developed in and today ecologists sit on advisory and sustain the biosphere.
Mexico in 1991, and then in boards, influencing both corporate
Agenda 21, an action plan adopted and government policies.
in 1992 at the United Nations Earth Despite such improvements, 2013, build on the work of the SBI.
Summit in Rio de Janeiro, Brazil. Lubchenco still believes that the They hope to effect greater change
Since 1991, the SBI and its report changes that have been made have in the next two decades, so that
have influenced a generation of not kept pace with the growing sustainable development can
ecologists, opening up new avenues dangers the planet faces. New satisfy humans’ current needs
of funding and collaboration, campaigns such as the ESA’s Earth without compromising the needs
forming committees, putting on Stewardship Initiative, created in of future generations. ■
Jane Lubchenco An acclaimed environmental Oceanic and Atmospheric

scientist and marine ecologist, Administration (NOAA). She
Jane Lubchenco grew up in was the first female and the first
Denver, Colorado. She earned a marine ecologist head of NOAA.
bachelor’s degree in biology at In 2011 Lubchenco oversaw the
Colorado College, and a Master’s creation of Weather-Ready Nation,
in zoology. She got her Ph.D. in a project to prepare the public in
marine ecology at Harvard. case of extreme weather.
She researches the interaction
between humans and the Key works
environment, with an emphasis
on biodiversity, climate change, 1998 “Entering the century of
and oceanic sustainability. the environment: a new social
From 2009 to 2013, she served contract for science,” Science
as Under Secretary of Commerce 2017 “Delivering on science’s
for Oceans and Atmosphere, and social contract,” Michigan
Administrator of the National Journal of Sustainability
324
WE ARE
PLAYING DICE
WITH THE NATURAL
ENVIRONMENT
THE ECONOMIC IMPACT OF CLIMATE CHANGE
C
limatology is an uncertain habitat destruction, shifts in
IN CONTEXT science. Future projections growing seasons, and enforced
will change, based on human migration.
KEY FIGURE
human actions and new technology,
William Nordhaus (1941–)
as well as natural cycles. However, Counting the cost
BEFORE it is vitally important to assess The social cost of carbon (SCC) is
1993 In Reflections on the the financial impacts of climate a monetary estimate of the damage
Economics of Climate Change, change. Once potential costs are to human society caused by every
William Nordhaus summarizes understood, we can explore ways additional tonne of carbon dioxide
the issues surrounding climate in which to mitigate its direct released into the atmosphere.
change and the economy, impacts. It is necessary to consider
highlighting uncertainties not only the direct costs—such as
Protesters in Lamu, Kenya,
and potential solutions. damage to property from flooding in 2018, opposing the construction
or fire—but also the costs of a coal-fired power plant. Growing
AFTER associated with broader effects, awareness of ecological damage has
2008 In Common Wealth: such as a decline in biodiversity, seen an increase in public disapproval.
Economics for a Crowded
Planet, Jeffrey Sachs argues
that although humanity faces
daunting economic crises—
including that of climate
change—we have the
knowledge to address them.
2013 The Climate Casino:
Risk, Uncertainty, and
Economics for a Warming
World, by William Nordhaus,
explains how global warming
relates to the world’s economy,
and provides ideas for
reducing its impact.

See also: Renewable energy 300–305 ■ Man and the Biosphere Programme
310–311 ■ Halting climate change 316–321
These damages include reductions emissions. The model divides the

in agricultural productivity, harm to world into distinct regions for its
infrastructure, energy costs, and analysis. It predicts that the
impacts on human health. The SCC combined SCC in 2055 will be
provides a starting point for energy between $40–$188 per ton ($44 and
policy. For example, if the SCC is $207 per tonne) of carbon dioxide
factored into proposals for a new released, depending on the rate of
power plant, the cost of building it warming and the mitigation
becomes much higher. This may policies enacted.
also make the cost of alternative Economic models incorporate
forms of energy—such as solar or assumptions, such as the discount William Nordhaus
wind power—more financially rate. Discount rates prioritize the
viable. However, it is extremely present over the future, because the Born in New Mexico in 1941,
difficult to calculate the SCC. future cannot be predicted perfectly. Nordhaus is a leader in the
The rate is selected based on how field of the economics of
Forecast models the balance between present and climate change. He stumbled
Economists use several models in future priorities is weighted. Higher upon this field of research
order to calculate the SCC. In 1999, discount rates indicate that future through sharing an office with
William Nordhaus developed RICE populations will be wealthier, and a climatologist. Nordhaus’s
(Regional Integrated Climate- prepared to deal with climate economic theories—the
Economy model)—a variant of his change. Lower discount rates DICE and RICE models—are
own preceding DICE (Dynamic suggest that the disruption caused widely used to analyze policy
decisions. Nordhaus is
Integrated Climate-Economy model), by climate change will make people
principally concerned with
which weighed the costs and in the future poorer than we are
placing a realistic price on
benefits of slowing down global today. Nordhaus suggests a 3 carbon. Today, the social cost
warming. The RICE model percent discount rate, meaning that of carbon is generally agreed
integrates carbon emissions, carbon if the monetary damages from to be around $40 per ton, but
concentrations in the atmosphere, climate change will be $5 trillion in Nordhaus’s models show that
climate change, damages, and the year 2100, we could invest $382 it should be higher to account
controls that are in place to reduce billion today to avoid it. ■ for the impacts of climate
change. Nordhaus is Sterling
Professor of Economics at Yale
Analyzing the costs of reducing carbon dioxide University, and serves on the
Congressional Budget Office
Panel of Economic Experts
Marginal Financial and the Brooking Panel on
benefits cost Economic Activity. In 2018,
THE COST OF REDUCING CO 2
Noordhaus was awarded the

Nobel Prize in economics.
The cost of reducing
CO2 increases in line
with the quantity, but Key works
this is offset by the
benefits gained. The 1994 Managing the Global
lines intersect at the Commons: The Economics
point of equilibrium, of Climate Change
where maximum 2000 Warming the World:
benefits are achieved Economic Models of Global
at the lowest cost. Warming
QUANTITY OF C0 2
326
MONOCULTURES AND
MONOPOLIES ARE
DESTROYING THE
HARVEST
SEED DIVERSITY
OF SEED
IN CONTEXT
KEY FIGURE
Vandana Shiva (1952–)
BEFORE
1966 A new high-yielding
strain of rice known as IR8
leads to a big increase in
production in several rice-
growing countries. First
developed in the Philippines,
it is also called “miracle rice.”
I
AFTER n 1987, Indian environmental Californian rice production is high
1994 The World Trade campaigner Vandana Shiva yield but there are problems with soil
launched a movement to protect salinity. Although salt tolerance can be
Organization introduces the genetically introduced, traditional rice
Trade Related Aspects of native seed diversity in response varieties can be naturally salt-resistant.
Intellectual Property Rights to changes in agriculture and food
(TRIPS) agreement. production. She founded Navdanya,
a nongovernmental organization, For example, a grass in the genus
2004 After protests by farmers to protect agricultural biodiversity Oryza was first cultivated for rice in
who developed the crop, the from the combined threat of genetic Asia between 8,200 and 13,500 years
Monsanto company’s patent engineering and patents. ago; today, there are more than
on an Indian strain of wheat 40,000 varieties of this rice in
known as Nap Hal is revoked. Agro-biodiversity existence. Intrinsic to agro-
Agricultural biodiversity (also known biodiversity are the many non-
2012 Indian initiative as agro-biodiversity) has resulted harvested species that support
Navdanya International from the selective breeding, over production. These include
launches its worldwide Seed thousands of years, of plants and microorganisms in the soil, species
Freedom campaign to protect animals taken from the wild. These that feed on pests, and pollinators.
food sovereignty and safety. practices led to the extraordinary Through the ages, the skills and
genetic diversity of different breeds knowledge of millions of farmers
of crops and domesticated animals. have shaped this biodiversity.

See also: Human activity and biodiversity 92–95 ■ Pesticides 242–247
■ Humankind’s dominance over nature 296 ■ Ecosystem services 328–329
new strains are patented by the

companies that created them.
Trade deals impose regulations
on who can use what. These work
against small-scale farmers but
favor the powerful agricultural
Seed patents threaten
corporations that produce the seed.
the very survival and
freedom of peasants … Seed sovereignty
and farmers … Shiva argues that rural farms are
Vandana Shiva threatened if the appropriate seed Vandana Shiva
is no longer available. Traditionally,
most small-scale farmers routinely Environmental campaigner
save their seed from one harvest to Vandana Shiva was born in
the next. Now, when farmers buy in northern India. Her mother
seed—especially if it is genetically was a farmer, and her father
modified—they often have to agree a forester. She studied in India
From the late 1960s, a technology not to save it. Having to buy seed and Canada, obtaining a
transfer to the developing world from a company every year can doctorate in the philosophy
included high-yield varieties leave them worse off financially. of physics. After returning
of cereals in association with Shiva criticized the practice of to India, in 1982 she founded
chemical fertilizers, pesticides, corporations patenting seed varieties the Research Foundation for
Science, Technology, and
and herbicides, mechanization, as “biopiracy” and set up Navdanya
Ecology. Following the Bhopal
and more efficient irrigation. to support “seed sovereignty.”
pesticide plant disaster in
Known as the “Green Revolution,” It campaigns for agro-biodiversity 1984, her interest in agriculture
this transformation shifted the via a network of seed-keepers and grew and three years later she
focus of agriculture in the developing organic producers and has helped founded Navdanya to protect
world away from biodiversity to found more than 100 community biodiversity and native seeds.
higher crop yields. New Green seed banks, effectively gene banks, Shiva campaigns against the
Revolution crops such as “miracle where seeds of crops and rare plant World Trade Organization’s
rice” (IR8) boosted production, but species are stored for future use. ■ Trade Related Aspects of
there was a downside. As more Intellectual Property Rights
emphasis was placed on fewer (TRIPS) agreement, which
productive strains, the genetic base broadens patents to include
of traditional seed varieties for plants and animals. TIME
grains, potatoes, fruits, vegetables, magazine hailed Vandana
and cotton declined. Shiva as an Environmental
Hero in 2003.
The United Nations Food and
Agriculture Organization estimates
Key works
that 75 percent of crop biodiversity
has been lost from the world’s fields. 1989 The Violence of the Green
Some environmentalists have Revolution
argued that traditional varieties are 2000 Stolen Harvest:
more compatible with local farming The Hijacking of the Global
Fertilizers have hugely increased
conditions, cheaper for farmers to food grain production in India—whose Food Supply
use, and more environmentally population of 1.3 billion people makes 2013 Making Peace with
sustainable than new, high-yield food security paramount—but the the Earth
varieties. Additionally, many of the chemicals also destroy soil fertility.
328
NATURAL ECOSYSTEMS
AND THEIR SPECIES HELP
SUSTAIN AND FULFIL
HUMAN LIFE
ECOSYSTEM SERVICES
IN CONTEXT
KEY FIGURE
Gretchen Daily (1964–)
BEFORE
c.400 bce Greek philosopher
Plato is aware of the human
impact on nature, noting that
deforestation can cause soil to
erode and springs to run dry.
1973 German statistician and
economist E.F. Schumacher
coins the term “natural capital”
in his book Small is Beautiful.
T
he benefits that humans As a sacred mountain, Mount Fuji
AFTER derive from ecosystems supplies a cultural ecosystem service
1998 The UN Environment are referred to by ecologists for the people of Japan, while the
program, NASA, and the surrounding rich volcanic soil provides
as ecosystem services. Some of the
World Bank release a study a service to the local tea plantations.
natural processes most important
on how protecting the planet to the continuation of human life
serves human needs. can be classified as ecosystem Although the idea that humans
services, such as pollination of benefit from nature has a long
2008 A study at the University
crops, decomposition of waste, history, it was not until the 1970s
of California, Berkeley, shows
and the availability of clean that the balance between nature
that ecological destruction by drinking water. Ecologists argue and human needs came to the
the world’s richest countries that because the enormous forefront of ecological debate. The
means they owe the world’s contributions of ecosystem services term “ecosystem services” first
poorest countries more than to human life are not readily appeared in the mid-1980s, and in
the developing world’s debt. quantifiable, humans drastically 1997 the concept was developed
undervalue these services while in two key articles: “Ecosystem
exploiting the natural world’s Services: Benefits Supplied to
resources for profit. Human Societies by Natural

See also: Human activity and biodiversity 92–95 ■ Ecological resilience 150–151
■ The Gaia hypothesis 214–217 ■ Human devastation of Earth 299
nature’s ability to control pests—

as opposed to humans’ use of
pesticides—and the atmosphere’s
capacity to clean itself naturally, as
well as the control of weather
If current trends hazards through natural buffers
continue, humanity will such as wetlands and mangrove
dramatically alter virtually forests. Pollination is another
all of Earth’s remaining important regulating service, one
natural ecosystems that is endangered by the global
within a few decades. decline of pollinators such as bees.
Gretchen Daily
Gretchen Daily Cultural services involve the ways
that humans assign cultural or Born in 1964 in Washington,
spiritual significance to elements of D.C., Gretchen Daily
ecosystems such as sacred trees, developed a passion for
animals, rivers, and mountains. ecology at a young age. After
The esthetic or recreational value her family moved to West
of a natural landscape is another Germany in 1977, she
Ecosystems,” edited by Gretchen type of cultural service. witnessed a national crisis
Daily, and “The Value of the World’s At its heart, the concept of over acid rain, and saw people
Ecosystem Services and Natural ecosystem services allows humans protesting in the streets over
Capital,” edited by American to see how inextricably connected environmental degradation.
ecological economist Robert they are to nature, and how without Daily earned two degrees and
Costanza. In 2001, UN Secretary the natural world human existence then her Ph.D. in biology at
Stanford University, where she
General Kofi Annan launched the would be impossible. Ecologists
is now the Bing Professor of
Millennium Ecosystem Assessment use the concept to illuminate Environmental Science.
(MEA), which helped popularize the how precious these systems are Daily studies biodiversity
concept of ecosystem services in for basic life conditions and to within the framework of
2005, when they published a wide- convince industries, businesses, “countryside biogeography,”
ranging appraisal of how humans and governments of the necessity or the portions of nature that
impact the environment. for ecological preservation. ■ have not been used for human
development, but whose
The four types of service ecosystems are still impacted
The MEA’s 2005 report detailed four by human activity. She is a
categories of ecosystem services: cofounder of the Natural
supporting, provisioning, regulating, Capital Project, which aims to
and cultural. Supporting services, incorporate environmentalism
Plans to protect air into business practices and
such as soil formation and water and water, wilderness and public policy.
purification, allow for the existence
wildlife are in fact plans
of all other services. Provisioning
services consist of freshwater;
to protect man. Key works
food, such as crops and livestock;
Stewart Udall
American politician and 1997 Nature’s Services:
fibres, including wood, cotton, conservationist Societal Dependence on
and other materials used for human Natural Ecosystems
essentials such as building and 2002 The New Economy of
clothing; and natural medicines, Nature: The Quest to Make
and plants used in pharmaceuticals. Conservation Profitable
Regulating services include
330
WE ARE LIVING ON THIS

PLANET AS THOUGH
WE HAVE ANOTHER
ONE TO GO
WASTE DISPOSAL
TO
M
ore than 65,000 people all adding to the world’s garbage
IN CONTEXT from at least 180 nations heap. Traditionally waste had been
traveled to Johannesburg, burned or buried, both options now
KEY FIGURE
South Africa, in 2002 to attend the associated with toxic greenhouse
Paul Connett (1940–)
United Nations World Summit on gas emissions and, in the case of
BEFORE Sustainable Development. Its final landfills, the potential for poisoning
1970 The first Earth Day resolutions included a call to ground water. The answer to the
takes place in the US to raise minimize waste and maximize world’s growing waste heap had to
awareness of clean waste reuse and recycling, and to develop be found elsewhere.
disposal and recycling. “clean” waste disposal systems.
In the last decades of the 20th The recycling revolution
1988 The Resin Identification century, it had become clear that Recycling for reuse is not a new
Code is introduced in the US refuse was reaching unmanageable concept, but its use as a way
to encourage the recycling proportions. Industrialization, the of reducing mountains of public
of plastic goods. growth of large urban populations, waste that would otherwise go into
and increasing use of plastic were landfill has its origins in the 1960s
1992 At the Rio Earth and 1970s, when organizations
Summit, 105 heads of state such as Greenpeace made the
pledge their commitment public more aware of environmental
to sustainable development. issues. Recently, campaigners such
AFTER as Paul Connett, author of Zero
2010 The United Nations Waste (2013), have renewed the
Pollution is nothing but global call to reduce consumption,
launches its Global Partnership the resources we are not and reuse or recycle items, rather
on Waste Management to harvesting. We allow them than discard them.
promote resource conservation to be dispersed because we’ve Since the 1970s, many US states
and efficiency. been ignorant of their value. and most European countries,
2012 Goals outlined at the R. Buckminster Fuller as well as Canada, Australia, and
UN Conference on Sustainable American inventor and architect New Zealand, have introduced
Development include waste curbside collections of recyclable
reduction and eco-friendly items sorted into bins. Sweden has
production methods. been especially active. In 1975,
Swedes recycled only 38 percent
of their rubbish, but today they lead

See also: Global warming 202–203 ■ Pollution 230–235 ■ Urban sprawl 282–283
■ A plastic wasteland 284–285 ■ Renewable energy 300–305
Methane from landfill
After carbon dioxide, methane
is the most critical greenhouse
gas. Although its atmospheric
concentrations are lower than
Refuse plastic CO2, methane is 25 times more
bags and excessive Rethink as you shop. powerful at trapping heat in
packaging. Buy products Do you really need what the atmosphere. Atmospheric
in large containers or you are buying? methane comes from various
without packaging. natural sources, including the
decay of vegetation in habitats
such as bogs and wetlands,
but also from livestock rearing,
from the use of fossil fuels,
Individual actions can and from the decomposition
reduce waste—households of trash in landfill sites.
in the developed world add a In many places, including
tonne of waste to landfill the UK and US, a number of
each year. landfill sites are now trapping
and collecting methane
to produce energy. Landfill
gas contains up to 60 percent
methane, depending on the
Reuse what you Recycle what composition of the waste and
can or pass it on can’t be used so that the age of the site. Vertical
to someone else it can be turned into and horizontal pipelines are
who can use it. new products. placed through the landfill
to collect the methane, which
is then processed and filtered.
Most of it is used to generate
the world, recycling 99 percent of silver, copper, and palladium used electricity, but it may also be
household waste. About 50 percent in circuit boards. It has been shown used in industry. After further
of this waste is burned in recycling that “mining” landfill sites to processing, it can be turned
plants that generate heat for the extract the metals can be more into fuel for vehicles, too.
nation’s homes. Sweden also cost-effective than mining natural
imports waste from other countries mineral deposits. However, e-waste
to process in its 32 incineration also includes toxic metals, such
plants. In 2015, it imported some as cadmium, lead, and mercury.
2.5 million tons (2.3 million tonnes) In countries that both generate and
of waste from Norway, the UK, import e-waste, landfill scavenging
Ireland, and other nations. for metals can be polluting. While
Europe now has an e-waste
“Mining” electronics reprocessing industry, relatively few
The fastest-growing type of waste efficient schemes exist elsewhere.
is discarded electronics. E-waste There are many new initiatives,
from mobile phones, computer hard but the world is still very far from
drives, TVs, and other electrical Connett’s zero waste ideal. A huge
goods reached almost 46 million challenge remains for individuals Methane is extracted at the
tons (42 million tonnes) in 2014— and governments: cut consumption Payatas landfill, Manila—the first
in the Philippines to have the gas
almost 25 percent more than in and recycle global refuse that will converted to energy, as part of a
2010. E-waste often contains soon reach 21/4 billion tons (2 billion United Nations program.
precious metals, such as the gold, tonnes) a year. ■
DIRECTO
RY
334
DIRECTORY
I
n addition to the scientists covered in the preceding chapters of this
book, many other men and women have made significant contributions
to the development of ecology. They have ranked among the greatest
scientific thinkers of their time. Some have excelled in academia, while
some came from other walks of life but pioneered new approaches to
advance. Still more have been formidable campaigners. Although they
worked in a range of disciplines, all have contributed to our understanding
of Earth’s biosphere, how it has evolved, and humanity’s place in it.
Crucially, their work continues to show what needs to be done to preserve
the natural world and to protect Earth from the destructive consequences
of human behavior.
undescribed species and used yarn notably in Canadian Wild Flowers (1865)
SAMUEL DE CHAMPLAIN to “band” birds—meaning he tied it to and Studies of Plant Life in Canada
1574–1635 their legs, allowing him to identify (1885). Her many albums of plant
individual bird—to find out more about collections are housed in the National
A French explorer, cartographer, soldier, their movements. Herbarium of Canada, at the Canadian
and naturalist, Champlain explored and See also: Animal ecology 106–113 Museum of Nature in Ottawa.
mapped much of Canada. He founded See also: Endangered habitats 236–239
the city of Quebec and established
the colony of New France. As a sharp MARY ANNING
observer and chronicler, he documented 1799–1847 KARL AUGUST MÖBIUS
animals and made notes about plants, 1825–1908
including details of leaves, fruits, and In 2010, the Royal Society named
nuts, and inquired about how the Anning as one of the 10 British women A German pioneer, Möbius was primarily
Native American people used them. who have most influenced the history of interested in the ecology of marine
See also: Classification of living things science. She found fame as a fossil ecosystems. After studying at the
82–83 collector and paleontologist, and her Natural History Museum of Berlin,
extraordinary fossil finds, from Jurassic and earning a Ph.D. at the University of
strata in the cliffs of the Dorset coast, Halle, he opened a seawater aquarium
JAMES AUDUBON included the first correctly described in Hamburg in 1863. While a professor of
1785–1851 ichthyosaur, two relatively complete zoology at the University of Kiel, his work
plesiosaurs, and the first pterosaur from on the viability of commercial oyster
The pioneer of North American outside Germany. Her finds helped production in the Bay of Kiel led him
ornithology, Audubon grew up in Haiti change views about Earth’s history, to recognize the various dependent
and France before emigrating to the providing strong evidence for extinction. relationships between organisms in
US in 1803. He developed an interest See also: Mass extinctions 218–223 the oyster bank ecosystem.
in nature, especially birds, and was See also: The ecosystem 134–137
a talented artist. His artistic technique
was unusual: after shooting a bird, CATHERINE PARR TRAILL
he held it in a “natural pose,” using fine 1802–1899 ERNST HAECKEL
wire, and painted it with a backdrop of 1834–1919
the bird’s natural habitat. Between 1827 A botanist and prolific author, Traill
and 1838 he published The Birds was born in the UK and emigrated to Haeckel was a biologist, physician,
of America in a series of installments. what is now Ontario, Canada, after she and artist who popularized Charles
It included 435 colored prints of 497 married in 1832. There, she wrote about Darwin’s ideas in Germany (while also
species, six of which are now extinct. life as a settler in Canada. She also rejecting many of them) and introduced
Audubon also discovered 25 previously wrote about the natural environment, the word “ecology” in 1866. Born in
DIRECTORY 335
Potsdam, he studied at several went on to pioneer phytosociology, the

universities before becoming a zoology study of natural plant communities, first MARJORY STONEMAN DOUGLAS
professor at the University of Jena in using the term in 1896. In the 1920s 1890–1998
1861. Haeckel was the first biologist Paczoski established the world’s first
to propose the kingdom Protista—for institute of phytosociology, at the A formidable campaigner for the
organisms that are neither animal nor University of Poznan, where he was protection of the Florida Everglades,
vegetable—and he researched and professor of plant systematics. An Douglas was also a successful journalist
painstakingly recorded tiny deep- accomplished botanist, he published and author, suffragist, and campaigner
sea protozoans called radiolaria. works on central European flora, for civil rights. Her 1947 book The
See also: Evolution by natural selection including that of the Białowieza Forest, Everglades: River of Grass was
24–31 which he managed as a national park. influential in building an appreciation
See also: Organisms and their of the Florida wetlands, and in 1969 she
environment 166 founded the Friends of the Everglades
WILLIAM BLAKE RICHMOND to defend the area from draining for
1842–1921 development. Douglas remained active
JACK MINER well into her second century, and at
Best known as a British artist, sculptor, 1865–1944 the age of 103 she was awarded the
and designer of stained glass and Presidential Medal of Freedom.
mosaics, London-born Richmond Also known as “Wild Goose Jack,” Miner See also: Citizen science 178–183
became an environmental activist moved with his family from the US to
after having to endure the poor light Canada in 1878. He was illiterate until
and smoky air produced by London’s the age of 33 but embarked on local BARBARA MCCLINTOCK
winter coal fires. In 1898 he founded conservation projects, such as building 1902–1992
the Coal Smoke Abatement Society winter feeding stations for Bobwhites.
(CSAS) to lobby politicians for clean He was one of the first people in North In 1983 McClintock became the first
air. The CSAS was instrumental in the America to put aluminum bands on solo woman to win the Nobel Prize in
introduction of the UK’s Public Health birds’ legs to track their movements. A Physiology or Medicine, and the first
(Smoke Abatement) Act in 1926 and duck banded by him, and later seen American woman to win any unshared
the Clean Air Act in 1956. in South Carolina, was the first banding Nobel Prize. The award recognized her
See also: Pollution 230–235 recovery made in North America. Miner discovery—more than 30 years before—
is thought to have banded more than of transposable genetic elements, or
90,000 wildfowl, helping establish “jumping genes,” which sometimes
THEODORE ROOSEVELT a huge database of migration routes. create or reverse mutations. As a
1858–1919 See also: Citizen science 178–183 cytogeneticist concerned with how
chromosomes relate to cell behavior, she
To deal with severe childhood asthma, also discovered the first genetic map
Roosevelt became an active sportsman JAMES BERNARD HARKIN for corn—linking physical traits with
and outdoorsman, developing a lifelong 1875–1955 regions of the chromosome—and the
passion for nature. When, in 1900, he mechanism by which chromosomes
stood as William McKinley’s running Sometimes referred to as the “father exchange information.
mate in the US presidential election, he of Canadian national parks,” Harkin had See also: The role of DNA 34–37
did so on a ticket of peace, prosperity, a passion for politics and conservation.
and conservation. Roosevelt became In 1911, he was appointed the first
the 26th President when McKinley was commissioner of the Canadian National JACQUES COUSTEAU
assassinated in 1901, and went on to Parks Agency and oversaw the 1910–1997
establish the US Forest Service, five establishment of Point Pelee, Wood
new national parks, 51 bird reserves, Buffalo, Kootenay, Elk Island, Georgian French undersea explorer Cousteau was
and 150 national forests. Bay Islands, and Cape Breton Highlands well known as the presenter of several
See also: Deforestation 254–259 national parks. Harkin realized the documentaries on the aquatic world.
commercial value of the parks, and After inventing underwater breathing
his policy of encouraging road-building apparatus called the Aqua-Lung in
JÓSEF PACZOSKI to attract tourists was not universally 1943, he worked with the French Navy
1864–1942 popular. He was a prime mover behind to clear underwater mines after World
legislation to regulate the hunting War II. He later converted the Calypso,
Paczoski was a Polish ecologist, born of migrant birds in 1917. a former minesweeper, into a research
in what is now Ukraine. He studied See also: Endangered habitats 236–239, vessel from which he explored the
botany at the University of Kiev and Deforestation 254–259 oceans, writing several books and
336 DIRECTORY
making hours of television. The Australia. He graduated from the women toured the country, speaking
Calypso was badly damaged in 1996, University of Sydney with a degree at meetings to highlight the dangers
but Cousteau died suddenly in 1997 in botany and zoology in 1937, and then of the dump, which they feared could
before he could afford to replace it. studied at Harvard University, earning grow as foreign governments and
See also: A plastic wasteland 284–285 a Ph.D. for his work on termites. After corporations saw an opportunity to
World War II, he returned to Australia, dispose of their radioactive waste.
where he became the first head of See also: Pollution 230–235
PIERRE DANSEREAU the Commonwealth Scientific and
1911–2011 Industrial Research Organization’s
Division of Forest Research in 1976. EUGENIE CLARK
Dansereau was a French Canadian Particularly known for his work on 1922–2015
plant ecologist who pioneered the study myxomatosis and its use in controlling
of forest dynamics and is considered rabbit populations, Day published his Known as the “Shark Lady” for her
one of the “fathers of ecology.” Born in first paper in 1938, and his last—on research on shark behavior, Clark was a
Montreal, he gained his Ph.D. in plant moths—74 years later. Japanese-American marine ecologist
taxonomy at the University of Geneva See also: Thermoregulation in insects and a pioneer in the use of scuba diving
in 1939. He later helped set up the 126–127 ■ Invasive species 270–273 for scientific research—she undertook
Montreal Botanical Garden and many dives around Florida’s Cape Haze
wrote numerous papers on botany, Marine Laboratory, where she worked
biogeography, and the interaction of JUDITH WRIGHT alongside other female ecologists such
humans and the environment. In 1988 1915–2000 as Sylvia Earle. Clark made several
he was appointed Professor Emeritus key discoveries about sharks and fish,
at the University of Montreal, a post he Principally a poet, Wright was also and was a major advocate of marine
held until he retired, aged 93, in 2004. renowned in her native Australia for conservation. In 1955, she founded the
See also: Biogeography 200–201 campaigning on Aboriginal land rights Mote Marine Laboratory, which works
and environmental issues. She was to protect shark species, preserve coral
born in Armidale, New South Wales, reefs, and found sustainable fisheries.
MARY LEAKEY and studied at the University of Sydney, See also: Animal behavior 116–117
1913–1996 before publishing her first book of
poetry in 1946. Between 1967 and 1971,
London-born Mary Leakey, one of the along with artist John Busst and DAVID ATTENBOROUGH
world’s foremost paleoanthropologists, environmentalist Len Webb, she built 1926–
experienced her first archeological an alliance of conservation groups,
excavation at the age of 17, when she was trade unions, and concerned citizens to British naturalist and television
hired as an illustrator at a “dig” in Devon. fight Queensland state government’s producer Attenborough served as a
In 1937 she married paleoanthropologist plans to open up the Great Barrier Reef controller for the BBC before stepping
Louis Leakey, and the couple moved to to mining. The campaign, detailed in down to dedicate more time to writing
Africa to work in the Olduvai Gorge— her book The Coral Battleground (1977), and producing documentaries. He
a site rich in fossils—in what is now eventually succeeded. wrote and narrated a series of nature
Tanzania. In 1948, Mary found the fossil See also: The Green Movement 308–309 programs, notably the Life series,
skull of an 18-million-year-old ancestor of beginning with Life on Earth (1979).
apes and humans, Proconsul africanus. Attenborough’s work has been credited
More breakthroughs in understanding EILEEN WANI WINGFIELD with renewing public interest in nature
human ancestry followed, including the 1920–2014 and conservation in Great Britain.
discovery in 1960 of Homo habilis, a See also: A plastic wasteland 284–285
1.4–2.3-million-year-old hominid who As a young Aboriginal woman in
used stone tools. Australia, Wingfield herded cattle and
See also: Evolution by natural selection sheep with her father and sister. In the PETER H. KLOPFER
24–31 early 1980s she lay down in front of 1930–
bulldozers at Canegrass Swamp in
opposition to construction of the Berlin-born Klopfer is an ecologist
MAX DAY Olympic Dam uranium mine. Later, whose main area of interest is ethology,
1915–2017 Wingfield teamed up with Eileen studying animal behavior in a natural
Kampakuta Brown and other Aboriginal environment. His influential 1967 book
An ecologist and entomologist, Day elders to campaign against the An Introduction to Animal Behavior:
developed an interest in wildlife, government’s proposals to dump Ethology’s First Century acted as a
particularly insects, as a boy in nuclear waste in South Australia. The survey and synthesis of past and
DIRECTORY 337
present ethological theories. In 1968, determines where animals choose to and the author of books on nature and
he began teaching in the Department live. His 1963 paper on habitat selection the environment. He cofounded the
of Zoology at Duke University, North by prairie deer mice demonstrated that David Suzuki Foundation in 1990 to
Carolina, where he was instrumental instinct and experience both play a role investigate sustainable ways for people
in starting its primate center. in how the mice select their habitat. to live in harmony with the natural world.
See also: Animal behavior 116–117 See also: Ecological niches 50–51 See also: Environmental ethics 306–307
DIAN FOSSEY SYLVIA EARLE DANIEL B. BOTKIN

1932–1985 1935– 1937–
Most of what is known about the lives An American marine biologist, author, Botkin, a prominent American author
and social structures of wild mountain and conservationist, Earle is an expert and environmentalist, earned his Ph.D.
gorillas in Africa derives from the work on the impact of oil spills. In 1991, she in plant ecology in 1968 at Rutgers
of primatologist and conservationist assessed the damage caused by the University. He writes and speaks on all
Fossey. The daughter of a San Francisco destruction of Kuwaiti oil wells during the areas of the environment, from forest
fashion model, she graduated and Gulf War. Earle undertook similar work ecosystems to fish populations, and also
worked as an occupational therapist after the Exxon Valdez, Mega Borg, and advises agencies, corporations, and
before visiting Africa, where she met, Deepwater Horizon oil spills. In 2009, governments. After decades spent
and was inspired by, Mary and Louis Earle launched Mission Blue, which, by researching climate change, Botkin has
Leakey. In early 1967 Fossey founded 2018, had established nearly 100 marine questioned how far it is impacted by
the Karisoke Research Center in the protected areas around the world. human activity. He is a research scientist
Rwandan mountains, where she See also: Pollution 230–235 at the Marine Biological Laboratory, near
studied gorillas. Her best-selling 1983 Boston, and is involved in environmental
book about her experiences—Gorillas studies programs at several American
in the Mist—was later adapted for the ROBERT E. SHAW universities.
screen. Fossey was murdered at her 1936– See also: Halting climate change 316–321
camp in December 1985, probably
because of her anti-poaching stance. Shaw is an American pioneer
See also: Animal behavior 116–117 of ecological psychology, which looks at EILEEN KAMPAKUTA BROWN
how perception, action, communication, 1938–
learning, and evolution in humans
TOMOKO OHTA and animals are determined by the In the early 1990s, the Australian
1933– environment. In 1977 he coedited the government revealed plans to build a
book Perceiving, Acting, and Knowing: nuclear waste dump near Woomera, in
Ohta is a Japanese population Toward an Ecological Psychology, the South Australian desert. Together
geneticist who in 1973 proposed the which effectively launched this new with Eileen Wani Wingfield, Brown, an
revolutionary Nearly Neutral Theory, area of study. In 1981 Shaw was the Aboriginal elder, established a kungka
which included the idea that mutations founding president of the International tjuta (women’s council) in the town of
that are neither neutral nor harmful play Society for Ecological Psychology and Cooper Pedy to fight the plans. They
an important part in evolution. After is now an emeritis professor in the were aware of the birth defects, cancer,
graduating from the University of Department of Psychological Sciences and other health issues following the
Tokyo in 1956, Ohta worked on the at the University of Connecticut. British military’s nuclear tests in the
cytogenetics (how chromosomes relate See also: Using animal models to desert in the 1950s and 1960s, and
to cell behavior) of wheat and sugar understand human behavior 118–125 feared that radiation could seep into the
beet, and now works at Japan’s groundwater. The plans were abandoned
National Institute of Genetics. and Brown and Wingfield won the 2003
See also: The selfish gene 38–39 DAVID SUZUKI Goldman Environmental Prize.
1936– See also: Pollution 230–235
STANLEY C. WECKER Canadian scientist Suzuki earned a

1933–2010 Ph.D. in zoology from the University of LYNN MARGULIS
Chicago in 1961, and two years later 1938–2011
An American animal behaviorist, became a professor in the genetics
Wecker was an influential researcher department at the University of British American biologist Margulis attended
into animal population and community Columbia. Since the mid-1970s, he has Chicago University aged only 15 and
ecology, especially the study of what also been a TV and radio broadcaster gained her Ph.D. at the University
338 DIRECTORY
of California, Berkeley, in 1965. The next work on chaos theory. In his 1975 paper movement. In the early 1980s, he was
year, at Boston University, she proposed “Period Three Implies Chaos,” written one of the leaders of a successful
that cells within nuclei had evolved with Chinese mathematician Tien-Yien campaign to prevent the building of
as a result of the symbiotic merger Li, he argued that above a certain rate the Franklin Dam, which would have
of bacteria. This idea, although not of growth, population forecasts become destroyed key habitats. In 1996, Brown
generally accepted until the 1980s, totally unpredictable, a discovery with was elected to the Australian Senate as
transformed the understanding of major ecological implications. a Green Party representative. On
cell evolution. See also: Population viability analysis retirement in 2012, he set up the Bob
See also: The Gaia hypothesis 214–217 312–315 Brown Foundation to campaign for the
protection of Australian habitats.
See also: The water crisis 288–291
PAUL F. HOFFMAN IAN LOWE
1941– 1942–
BIRUTE GALDIKAS
Canadian scientist Paul Hoffman’s Lowe, an Australian environmentalist 1946–
discovery of “cap carbonates”—evidence who studied engineering and science at
for ancient glaciation in Precambrian the University of New South Wales and German-born anthropologist and
sedimentary rocks in Namibia—revived earned his Ph.D. in physics at the primatologist Galdikas has pioneered
the “Snowball Earth” hypothesis in University of York, advises the UN’s the study of orangutans in the wild.
climate change studies in 2000. The Intergovernmental Panel on Climate Along with Jane Goodall and Dian
term was first used by American Change. He is outspoken on the need Fossey, she was one of “The Trimates,”
geologist Joseph Kirschvink in 1992, for renewable energy, arguing that it chosen by Louis Leakey to study great
although there had been speculation is “quicker, less expensive, and less apes. Leakey persuaded her to support
since the late 19th century that Earth’s dangerous than nuclear power.” In the establishment of an orangutan
surface was almost entirely frozen more 1996, Lowe chaired the expert group research station in Borneo, to which she
than 650 million years ago. responsible for the first report on the moved in 1971. For more than 30 years,
See also: Ancient ice ages 198–199 state of Australia’s environment. Lowe Galdikas studied the great apes,
is now Emeritus Professor of Science, advocated protection for them and their
Technology, and Society at Griffith rain forest habitat, and undertook the
SIMON A. LEVIN University, Brisbane. rehabilitation of orphaned orangutans.
1941– See also: Renewable energy 300–305 See also: Animal behavior 116–117
■ Halting climate change 316–321
Levin, an American ecologist, specializes

in the use of sophisticated mathematical BRIAN A. MAURER
modeling, alongside field and lab AILA KETO 1954–2018
observation, to understand the workings 1943–
of ecosystems. He also researches the Maurer’s 1989 paper “Macroecology:
relationships between ecology and Keto spent much of her youth exploring The Division of Food and Space Among
economics. Levin earned a Ph.D. the Great Barrier Reef and surrounding Species on Continents”—written with
in mathematics from the University rain forests. She studied biochemistry James H. Brown—was the first clear
of Maryland in 1964 and taught at and went on to work at the University articulation of the idea that there is value
Cornell University from 1965 to 1992. of Queensland. In 1982, with her in studying ecological patterns and
After moving to Princeton, he was husband Keith, she founded the processes over large areas and long time
appointed director of the university’s Australian Rain forest Conservation frames. In his last years he researched
Center for BioComplexity, which Society, which did much to save the dynamics of the spread of exotic birds
investigates the mechanisms that Australia’s Wet Tropics area. and species diversity among mountain-
generate and maintain complexity See also: Biomes 206–209 dwelling mammals in North America.
in the living world. See also: Macroecology 185
See also: Predator–prey equations 44–49
BOB BROWN
1944– NANCY GRIMM
JAMES A. YORKE 1955–
1941– After studying medicine at the
University of Sydney, Brown practiced Based at Arizona State University,
An American mathematician and in Australia and the UK. He moved to Grimm is a climate change ecologist
physicist based at the University of Tasmania in 1972 and soon became and sustainability scientist, whose
Maryland, Yorke is best known for his involved in the environmental research concentrates on the
DIRECTORY 339
interaction of climate change, human such as the Hawaiian chain, where the
activity, and ecosystems. Her work has date of each island is already known SARAH HARDY
particularly focused on the movement with some accuracy. Most of her work 1974–
of water and chemicals through is focused on the evolution of spider
ecosystems. Grimm is a past president species. Gillespie is based at the Hardy is an American marine biologist
of the Ecological Society of America University of California, Berkeley, where and polar explorer who studies the effect
and a senior scientist on the US Global she runs the EvoLab, a research group on the environment of deep-ocean
Climate Change Research Program. that focuses on arthropods, such as mining. She argues that to protect
See also: Ecosystem services 328–329 spiders and insects. marine communities and biodiversity
See also: Thermoregulation in insects it is important to develop a systematic
126–127 ■ Island biogeography 144–149 approach to the zoning of the oceans—
TIM FLANNERY with deep-sea marine protection areas a
1956– priority. Hardy studied marine biology at
HARVEY LOCKE the University of California and earned
One of Australia’s most prominent 1959– her Ph.D. in oceanography at the
environmentalists, Flannery earned University of Hawaii in 2005.
a Ph.D. in 1984 from the University of Born in Calgary, Canada, Locke trained See also: A plastic wasteland 284–285
New South Wales for his work on and practiced as a lawyer before
kangaroo evolution. He later built switching to full-time conservation
a reputation as a mammalogist, work in 1999. He is committed to areas KATEY WALTER ANTHONY
discovering several new species, and of ecology known as large landscape 1976–
as an expert on climate change. He and connectivity conservation, which
was chief commissioner of the Climate involve the connection of all lands, Based at the University of Alaska,
Commission, an Australian government whether urban or wild, across a wide Walter Anthony is an aquatic
body, and champions renewable energy. network. Locke was a founder of the ecosystems ecologist specializing in
See also: Renewable energy 300–305 Yellowstone to Yukon Conservation polar environments. She has studied
Initiative, which campaigns to create carbon dioxide and methane emissions
a continuous wildlife corridor between from lakes in the North American
SUSAN KAMINSKYJ those two areas of North America. In Arctic. In 2017, she discovered that
1956– 2009, Locke also cofounded the Nature unusually large amounts of methane
Needs Half movement, which advocates were escaping from an Arctic lake,
From her laboratory at the University for the protection of half of Earth’s land where the gas was seeping into the
of Saskatchewan, Canada, Kaminskyj— and water area by 2050. Locke argues water from greater depths than
a cell biologist and mycologist—has that this policy is necessary to avoid previously discovered. If replicated
pioneered the use of fungi to clean a sixth mass extinction on Earth. elsewhere, such emissions from
oil-contaminated site, in a process See also: Mass extinctions 218–223 reserves deep in the permafrost could
known as bioremediation. Kaminskyj produce a dramatic increase in the
and her team found that when seeds amount of methane in the atmosphere.
are treated with a fungus named MAJORA CARTER See also: The Keeling Curve 240–241
TSTh20-1, plants can establish in the 1966–
substrate of such land and clean the
soil as they grow. When her dog led her through a AUTUMN PELTIER
See also: Ubiquity of mycorrhizae degraded brownfield site to the banks 2004–
104–105 ■ Pollution 230–235 of the Bronx River, in her native New
York City, Carter realized the potential Peltier, a member of the Wikwemikong
for the regeneration of this area. She First Nation who lives in Ontario,
ROSEMARY GILLESPIE won funding from the city council to Canada, is a campaigner for clean
1957– develop Hunts Point Riverside Park on drinking water, arguing that humanity
the site, providing a natural retreat and should treat water with greater respect.
Scottish-born Gillespie studied zoology river access for locals. Subsequently In 2018, at the age of 13, she was one
at the University of Edinburgh before her organization, Sustainable South of the youngest people ever to speak
moving to the US to earn her Ph.D. at Bronx (SSBx), advocated and won to the UN General Assembly. Here, she
the University of Tennessee. She is support for “green” urban renewal in advocated the policy that “No child
known particularly for investigations disadvantaged communities elsewhere should grow up not knowing what clean
into what drives biodiversity at species in New York. SSBx also campaigns to drinking water is, or never know what
level, concentrating her evolution improve air quality and food choices. running water is.”
research on “hotspot archipelagos” See also: The Green Movement 308–309 See also: The water crisis 286–291
340
GLOSSARY
Abiotic Nonliving; often used to Biogeography The study of how Citizen science Scientific research
refer to the nonliving components plants and animals are distributed carried out by amateurs, typically
of an ecosystem (such as climate geographically, and the changes involving large-scale data collection.
and temperature). to this distribution over time.
Climate change A shift in the
Abundance The number Biological community A world’s interconnected weather
of a given species within an collection of living organisms patterns; a gradual natural process
ecosystem; an abundant species within one location; when exacerbated by human actions.
is strongly represented within the combined with their environment,
wider population. they make an ecosystem. Climax A biological community
or ecosystem that has reached
Acid rain Any form of precipitation Biomass The total quantity of a a stable point, so that populations
with high levels of acidity, causing given organism within a habitat, of organisms will remain steady.
damage to the environment; may generally expressed as weight or This is the end result of succession,
occur naturally or as a result of volume. Also a type of fuel made in which the type of species and
human activity. from organic matter, usually burned population sizes that make up
to generate electricity. a community change over time.
Anthropogenic Originating in,
or influenced by, human activity. Biome An area of Earth that can Climax species A plant species
be classified according to the that will not change as long as its
Apex predator A predator that species of plant and animal life environment remains stable.
is not prey for any other species. within it.
Clutch size The number of eggs
Atmosphere The layer of gases Biosphere The layer of Earth laid in one birthing.
surrounding Earth. It also protects in which life can exist, situated
organisms from ultraviolet radition. between the atmosphere and Community ecology The study of
lithosphere; the sum of all how species interact within a given
Autotroph A producer; an ecosystems on the planet. geographical space.
organism that makes its own food
from sources such as light, water, Botany The scientific study Competitive exclusion
and chemicals in the air. of plant life. principle The idea that multiple
species reliant on exactly the same
Behavioral ecology The study of Carnivore An organism which resources cannot exist together
animal behavior and how ecological eats only meat. without one population rising and
pressures influence this. the other falling, as one will always
Catastrophism The theory that have an advantage over another.
Biodegradable Usually used changes in Earth’s crust were
in reference to waste products, caused by dramatic and unusual Coniferous Describes trees with
meaning something that can be events, as opposed to gradual seed cones which mostly do not
broken down by natural processes. change over time. shed their needlelike leaves
during winter.
Biodiversity The variety Cells The smallest structural and
of ecological life within a given biological unit that can survive Conservation The protection and
geographical area, encompassing on its own; the “building blocks” preservation of animal life, plant
variety between and within species. of all life on Earth. life, and natural resources.
GLOSSARY 341
Consumer A species that eats Endangered Describes a species the connections between them,
other organisms to obtain its whose population is so small that illustrating how communities
required nutrients; this term can it is at risk of dying out completely. interact on a wider scale to survive.
apply to any organism that is not
at the very bottom of the food chain Epidemiology The study of Fossil The remains of a prehistoric
how diseases spread through organism, preserved and solidified
Deciduous Describes trees that populations, and the impact this in sedimentary rock or amber.
shed their leaves in the fall. has on the wider ecosystem.
Fossil fuel Nonrenewable fuels
Decomposers Organisms, Ethology The scientific study formed over millions of years from
primarily bacteria and fungi, that of the evolution of animal behavior plant and animal remains.
break down dead organisms and as an adaptive trait, with a
waste matter to obtain energy. particular focus on observing Fracking A process by which oil
animals in their natural habitat. or gas can be extracted from the
Deforestation The cutting down ground. Fracking involves drilling
of a large area of trees, carried out Evolution The process by which down and injecting liquid into
for a range of purposes, including species change over time as traits the rock at a high pressure in
farming, industry, and construction. are passed down over generations. order to force the oil and gas to
the surface.
Detritivores Organisms that feed Extinction The permanent dying
on waste matter. out of an entire species. Fungi A group of organisms,
including mushrooms, that produce
Diatom Any of a large group of Extirpation Extinction of a spores and feed on organic matter.
microscopic algae that often play species on a local level—when Unlike plants, fungi do not utilize
an important role in stabilizing an a species dies out within a specific sunlight for growth.
ecosystem and facilitating the geographic area but still exists
existence of a range of life forms. elsewhere on the planet. Gene The most basic unit of
heredity; part of a DNA molecule
Diversity A measure of the variety Feedback loop The effect that one that transmits characteristics from
of species within a biological part of an ecosystem has on the a parent to its offspring.
community or ecosystem. rest, and how this change feeds
back into the system as a whole. Genome The complete set of an
DNA Deoxyribonucleic acid. A organism’s genes.
large molecule in the shape of a Fertilizers Substances, which
double helix that carries genetic can be either natural or chemical, Geology The scientific study
information in a chromosome. that are added to soil to increase of Earth’s physical formation and
its nutrient content and help plants structure. Geologists examine our
Ecology The scientific study of grow more successfully. planet’s history and the ongoing
the relationships between living processes that are acting upon it.
organisms and their environment. Fieldwork Studies undertaken
in the wild, rather than under Global warming A gradual
Ecosystem A community of controlled laboratory conditions. increase in the temperature of
organisms in a given environment Earth’s atmosphere caused by the
that interact with and affect Food chain A series of predators accumulation of greenhouse gases.
one another. and prey, in which each organism
is dependent on the preceding GMO Genetically Modified
Ecosystem services The benefits lifeform for food. Organism—any life form that has
humans receive from an ecosystem; been artificially and chemically
a term highlighting the importance Food web A collection of food altered by engineering techniques
of the environment to humanity. chains within an ecosystem and that modify its DNA.
342 GLOSSARY
Greenhouse effect The way in Irrigation The controlled Migration A large-scale movement
which gases in Earth’s atmosphere application of water to areas of of a species from one environment
trap heat. The buildup of these land, usually through the creation to another; often occurs seasonally.
gases leads to global warming. of channels, to help crops grow.
Monoculture Using land for the
Greenhouse gas Gases such as Keystone species A species cultivation or growth of only one
carbon dioxide and methane that that plays a centrally important type of plant or animal. This often
absorb energy reflected by Earth’s role in an ecosystem, often has damaging effects on the land,
surface, stopping it from escaping disproportionate to its biomass, as it can decrease its mineral value.
into space. and whose removal would alter
or endanger the entire ecosystem. Morphology The study of the
Green Movement A political external structure of organisms.
ideology that encourages a greater Kin selection An evolutionary
focus on the importance of the strategy whereby individuals Mutation A change of
environment, and asks people to pursue the best tactic for their structure within an organism’s
take action to prevent damage relatives’ survival, even at the DNA, which may result in a
to Earth’s natural habitats. expense of their own safety, well- genetic transformation giving
being, or reproduction. it uncharacteristic traits. One
Groundwater Water found below example of a mutation is
Earth’s surface, such as in spaces Mass extinction The widespread albinism, a lack of pigmentation.
in the soil, sand, or rock. and rapid dying out of an abnormally
large number—at least half—of Mutualism A situation in which
Habitat The area in which all species; this sharp change in two or more organisms depend
an organism naturally lives. biodiversity usually marks a shift on each other for survival.
to a new geological era in our
Herbivore An organism that eats planet’s history. Mycorrhizae Types of fungi
only plants. that grow among the roots of
Metabolism The chemical plants and exist in a symbiotic
Homeostasis The regulation processes that occur within the relationship with these plants.
of conditions within an organism, cells of an organism to keep it alive,
such as temperature, water, and such as the processes that enable Natural selection The process
carbon dioxide, to maintain a the digestion of food. by which characteristics that
stable internal state. increase an organism’s chances
Metacommunity A set of of reproducing are preferentially
Hypothesis An idea or assumption, independent communities that passed on.
used as the starting point for interact and are connected by
a theory, which is then tested the movement of some species Niche The specific space and
through scientific experimentation. between those communities. role that a species occupies within
an ecosystem.
Inheritance The passing on of Metapopulation A collection
genetic qualities and behavioral of smaller populations of a given Omnivore An organism that feeds
predispositions to offspring, species that are linked by the on both animals and plants.
through both genetic information movement of some individuals.
and parental nurture. Organism General term for
Microorganism An organism, any living thing, from single-cell
Invasive species A nonnative invisible to the human eye, bacteria to complex, multicellular
species that has been introduced that can only be seen with a life forms such as plants and animals.
to an ecosystem and spreads microscope, such as a bacterium,
rapidly, damaging the ecological virus, or fungus; also known as Ornithology A branch of biology
balance of the area. a microbe. that concerns the study of birds.
GLOSSARY 343
Overfishing The depletion of the Prey A species that is hunted Thermoregulation The internal
fish population in a given area as by another species. processes that occur within an
a result of fishing too intensively. organism to ensure it maintains
Primary producer Any organism a stable temperature, a function
Ozone layer Part of the upper that makes its own food from that is crucial for survival.
level of Earth’s atmosphere, with nonorganic sources, namely light
a high concentration of ozone (O3) and/or chemical compounds such Transmutation The process of
molecules; provides protection as carbon dioxide and sulfur, and evolutionary divergence by which
from ultraviolet radiation. thus sustains the animals that one species transforms into an
feed on it. entirely new one.
Paleontology The study of fossils
and biology of Earth’s geological Primary vegetation The Trophic cascade The impact
past. Paleobotany is the branch vegetation that has prevailed in a that the removal of a trophic level
studying plant fossils. given area since the start of its of a food chain with at least three
current climatic conditions. levels has on the wider ecosystem
Parasite An organism that lives on as a whole.
or in another organism, and obtains Recycling The process of
nutrients from its host. converting waste into new objects Trophic level The place of an
or materials, or burning it to organism within an ecosystem’s
Pesticides Chemicals used to kill generate energy. hierarchy; organisms that are
certain types of pest in order to on the same level of the food chain
protect cultivated plants. They can, Renewables Fuel sources that are on the same trophic level.
however, also kill nontarget species are not finite, such as solar power,
and damage the wider ecosystem. hydropower, and wind power. Tropics The region of Earth that
surrounds the equator, between the
Photosynthesis The process Species A group of organisms lines of the Tropic of Cancer and
by which plants and algae transfer capable of exchanging genes with the Tropic of Capricorn, and does
the Sun’s light energy into chemical one another through reproduction. not experience the same seasonal
energy as glucose, allowing it to be changes as the rest of Earth.
passed along the food chain. The Stochasticity Unpredictable
process absorbs carbon dioxide and fluctuations in environmental Urbanization The process which
releases oxygen. conditions that affect populations occurs when rural areas are built
and ecological processes. upon intensively, almost always
Physiology A branch of biology with negative consequences for
that focuses on the everyday Succession The process by which the natural environment.
functioning of organisms. a biological community evolves
over time, from a few simple species Urban sprawl The outward
Pollination The transfer of pollen to a complex ecoystem, through growth of a previously concentrated
from a male plant part to a female species’ impact on the environment. urbanized area, often with negative
one—by birds, insects, and other consequences for the environment.
animals, or by the wind—enabling Taxonomy The science of naming
fertilization and seed production. and classifying different organisms. Variation Differences within a
species, caused either by genetic
Pollution The introduction of Tectonic plates Pieces of Earth’s or environmental factors.
harmful contaminants to the crust and uppermost mantle that
natural environment, inducing gradually shift over time, causing Vascular plant A type of plant
changes to the atmosphere. seafloor spreading, continental with conductive tissue for the
drift, and mountains, rift valleys, movement of water and minerals
Predator A species that hunts volcanoes, and earthquakes at throughout, such as a fern or a
other animal species for food. plate boundaries. flowering plant.
344
INDEX
Numbers in bold refer autogenic ecosystem engineers 189 biosphere 95, 136, 153, 160, 197, 204–205, 215
to a topic’s main entry autotrophs 132 biosphere reserves 236, 310, 311
Avery, Oswald 19, 34 Man and the Biosphere Programme (MAB)
axolotls 283, 283 310–311
A Sustainable Biosphere Initiative (SBI)

322–323
B
bipedalism 72
birds
bird counts 180–181, 182, 183
acacia trees 57–58, 57
birdsong 235
acid rain 93, 222, 229, 234, 248–249
eggs 114–115
adaptation 72
Bacon, Francis 294, 296, 296 light pollution damage 253
see also ecophysiology; natural selection
Bacon, Roger 84 migration 180, 180, 199, 278
Agassiz, Louis 196, 198–199
bacteria 30, 31, 31, 68, 69, 84, 85, 90, 100, social behavior 189
Age of Discovery 80, 296
102, 103, 136, 139, 164 bison 143
Age of Enlightenment 18
Bak, Per 184 black widow spiders 39, 39
aggression 124–125, 124
Aguado, Catalina 181 balsam fir 151 Blackburn, Tim 185
air pollution 93, 95, 232, 233–234, 233, 248 Baltimore Orioles 199 blood-suckers 127
Al-Jahiz 108, 130, 132 Banks, Jonathan 276 Blue Tits 114, 114
albinism 30 bark beetles 277 Blue-footed Boobies 115, 115
algae 132, 151, 217 Barlow, Maude 288, 289, 289, 291 Blue-gray Gnatcatchers 176–177, 176
algal blooms 166, 269 barnacles 54, 55, 55, 62, 63 bluebirds 111
alleles 30 Barro Colorado Island 157, 157 Bodenheimer, Frederick 112
allogenic ecosystem engineers 189 Bartram, William 297 boneworms 139
altruistic behavior 19, 29, 38, 39, 125 Bateson, William 29 bonobos 120, 123, 125, 125
Alvarez, Luis and Walter 22, 221, 221 bats 54, 67, 155 Bonpland, Aimé 162
Amazon Basin 97, 259 Bazalgette, Joseph 233 Botkin, Daniel B. 337
amphibian viruses 280 bears 51, 51, 72, 109, 109, 191, 313, 314, 314 Boyle, Robert 85
Anderson, Roy 68, 70–71 beavers 65, 65, 111, 188, 189 Bradley, Richard 130, 132
Andrews Forest, Oregon 153, 153 bees 29, 29, 38, 39, 66, 85, 100, 101, 126, 127, Brown, Bob 338
animal behavior see ethology 278, 279 Brown, Eileen Kampakuta 337
animal ecology 106–113 behavioral ecology 154–155 Brown, James H. 131, 146, 148, 185, 338
behavioral ecology 154–155 Beklemishe, Vladimir 204 Bruckner, John 132, 133
see also ecological niches; food chains; food webs Beneden, Pierre-Joseph van 56, 58 Brugger, Ken 181
Anning, Mary 334 benthic communities 142 Buckland, William 199
Anthony, Katey Walter 339 Biblical flood 198, 198, 199 budworm 151, 151
Anthropocene epoch 322 big ecology 153 buffalo 95, 110
anthropogenic biomes 95 Big Garden Birdwatch 182 Buffon, Comte du 20, 23, 26
antibiotics 103 bioaccumulation 94 bullfrogs 280
antibodies 103 biodiversity 63, 81, 90–97, 131, 137, 149, 235, bushmeat 124
antiseptics 103 237, 258, 322 butterflies 127, 181–182, 277, 279, 313
ants 48, 57–58, 57, 94, 142 agricultural 326–327
aphids 49, 58, 224 and ecosystem functioning 156–157
C
aquaculture 269 effects of human activity on 93
aquifers 289, 290, 291 hotspots 96–97
Aral Sea 288, 290, 290 key threats to 93–95
archaea 91 loss 156, 157
Arditi-Ginzburg equations 46 neutral theory of 152
argan trees 311 biofuels 289 C:N:P ratios 74, 75
Aristotle 42, 80, 82–83, 83, 100, 130, 166, 296 “biogenic” rocks 30 cactus 173
Arrhenius, Olaf 185 biogeography 94, 130–131, 162–163, 166, 197, Caldeira, Ken 281
Arrhenius, Svante 202–203, 240, 318–319 200–201, 209 Callendar, Guy Stewart 240
atmosphere 197, 204, 215 biological species concept 88–89 camels 73
Attenborough, David 93, 167, 336 biomass 62, 112, 113, 238, 305 camouflage 83
Audubon, James 181, 334 biomass energy 305 cancer research 75
Audubon Society 182 biomes 95, 135, 173, 197, 206–209, 209 cane toads 273, 273
INDEX 345
carbon 74, 75 Connett, Paul 330 barcoding 37

carbon dioxide (CO2) 65, 136, 228, 259, 281, conservation 124, 142, 236, 239, 267, 295, junk DNA 123
302–303, 305, 318, 321 307 mitochondrial DNA analysis 81
Keeling Curve 240–241, 241 see also environmentalism mutations 19, 30, 30, 31, 36–37
social cost of carbon (SCC) 324–325 continental drift 196, 212–213 sequencing 37, 89
carbon sinks 264 convergent evolution 209 Dobzhansky, Theodosius 29–30, 31, 96
carnivores 109, 133, 141 Cooke, Wells 180 dodos 299
Carson, Rachel 138, 139, 229, 244–245, 244, cooling ponds 137 dogs 89, 101, 116
247, 299, 306 cooperative behavior 124 Dolly the sheep 34
Carter, Majora 339 coral bleaching 207, 238 dolphins 97, 285, 291
Caswell, Hal 152 coral reefs 135, 152, 189, 193, 203, 205, 207, donkeys 89
Celsius, Anders 87 207, 238 Dorling, Danny 250
Central Park, Manhattan, New York 149 Cousteau, Jacques 335–336 Douglas, Marjory Stoneman 335
cephalopods 222, 223 Cowles, Henry Chandler 160, 170, 172, 174 dragonflies 43, 51, 76–77, 77, 109, 111, 189
CFCs 229, 260, 261 crabs 142 droughts 70, 288, 318
Chambers, Robert 20 Crawford, David 253 Drude, Oscar 172
chameleons 185 creationism 18, 20, 22, 28, 196 dunes 170, 172
Champlain, Samuel de 334 Crick, Francis 19, 32, 34–35, 35 dung beetles 126, 127
Chapman, Frank 181 crocodiles 42, 222 Dutch elm disease 70
Chargaff, Erwin 182 Croll, James 224
Charpentier, Jean de 198 Cronquist, Arthur 175
cheetahs 47 crows 67
chemotrophs 133
chestnut blight 175
Chicxulub Crater 220, 221, 237
chimpanzees 101, 120, 121, 121, 122, 122,
cuckoos 199
Curtis, John 161, 174, 175
Cuvier, Georges 18, 22
cyanobacteria 161, 189, 205
E
123–125, 123, 124 cystic fibrosis 37
E. coli 31, 31
cholera 69–70, 233
eagles 133, 229, 235
Christmas Bird Count (CBC) 180–181
D
chromosomes 123 Earle, Sylvia 337
Chutkan, Robynne 102 Earth Day 211, 211, 295, 306, 308, 330
chytridomycosis 280 Earth Summits 153, 323, 330
citizen science 161, 178–183 earthworms 189
cladistics 86, 87, 90, 91 Easter Island 264, 264
classification 20, 37, 81, 82–83, 86–87, 90–91 Dachille, Frank 220 echidnas 209
Clements, Frederic 135, 138, 152, 160–161, 166, Daily, Gretchen 329, 329 echolocation 67
167, 168, 170, 172–73, 174, 175, 197, 206–207, Daisyworld 216 ecological drift 192
208, 210 damselflies 109 ecological equivalence 51
climate change 95, 109, 113, 185, 202–203, 207, D’Ancona, Umberto 46–47, 48 ecological footprint 322
223, 224, 225, 228–229, 267, 268–269, 276, 281, Dansereau, Pierre 336 ecological niches 22, 43, 50–51, 108, 110–112,
295 Darwin, Charles 18, 21, 22, 23, 26–28, 26, 29, 176, 192
climate change denial 320 32, 42, 56, 59, 72, 116, 120, 130, 133, 146, competitive exclusion principle 42–43, 52–53,
economic impact 324–25 150, 162, 167, 193, 200, 297 112
halting 316–21 Dawkins, Richard 19, 38–39, 39, 88, 116, 123, generalists and specialists 111
Intergovernmental Panel on Climate Change 154 guilds 152, 161, 176–177
(IPCC) 276, 295, 299, 319–320, 321 Day, Max 336 niche construction 188–189
and spring creep 276–279 DDT 229, 244–245, 246, 247 niche overlap 51, 111–112
climax communities 172–173, 174 “dead zone” (oceans) 269 niche partitioning 51, 112
cloning 34 decomposers 139, 141 ecological pyramids 112, 112
clownfish 59, 59 decoupling of interactions between species ecological resilience 131, 137, 150–151
cockatoos 201 278–279 ecological stoichiometry 43, 74–75
cod fisheries 266–267, 268 deer 49, 77, 97 ecological succession 170–171, 171, 172, 173
“cold-blooded” 126 deforestation 93, 97, 228, 237, 238, 250, ecophysiology 72–73
colobus monkeys 124 254–259, 264, 294 ecoregions 237–238
competition detritus feeders 133 ecosystem ecology 157
competitive exclusion principle 42–43, 52–53, developmental traps 279 ecosystem services 328–329
112 diatoms 112, 189 ecosystems 128–159
exploitation 53 dinosaurs 22, 222, 223 biomes 95, 135, 173, 197, 206–209, 209
food chains and webs 109 Diogenes 297 biotic and abiotic elements 135–136, 208
interference 53 disease categories 136
interspecific 53 ecological epidemiology 68–71 ecological epidemiology 68–71
intraspecific 53 infectious diseases 280 energy flow 134, 138–139
Connell, Joseph 43, 55, 62, 170 DNA 19, 26, 30, 32, 34–37, 38, 123 equilibrium 136, 137
346 INDEX
ecosystems cont. evolutionarily stable strategy (ESS) 131, 154–155 forests see deforestation; rain forests
evolutionarily stable strategy (ESS) 131, evolutionary theory 16–39 Fossey, Dian 336–337
154–155 coevolution 56, 59 fossil fuels 93, 203, 217, 225, 240, 263, 302,
experimental 157 convergent evolution 209 319
external disturbances 136–137 early theories 20–21 fossils 18, 20, 21, 22, 23, 26, 30–31, 105, 196,
feedback loops 136, 217, 224–225 heredity 18, 19, 20, 21, 26, 28, 32–33 212, 212, 213
guilds 161, 176–177 kin selection 19, 29, 39 Fourier, Joseph 203, 299
holistic concept 175, 210–211 natural selection 18–19, 21, 22, 24–31, 38, 66, 72, foxes 132, 225, 315
keystone species 43, 60–65, 109, 130, 142 81, 154 fracking 304
mutualism 42, 43, 56–59, 100, 104, 105, 157 Red Queen theory 46, 49 Frank, Albert 100, 104, 105
resilience 131, 137, 150–151 see also DNA; genetics Franklin, Rosalind 19, 35
techno-ecosystems 137 extinction 22, 81, 92–93, 95, 96, 143 Friends of the Earth 297, 308
see also biodiversity; competition; ecological current rate of 92–93 Frisch, Karl von 116, 123
niches; food chains; food webs; predators extinction risk assessment 312–315 Fritts, Charles 302
and prey extirpation 93, 95 frogs 109, 280
ecozones 209 Holocene extinction 223 fruit bats 155
ectotherms 126, 222 island species 147, 148 fruit flies 75, 164–165, 165
Eden Project, UK 137 K-Pg extinction event 221–222 fungal diseases 280
eggs mass extinctions 22, 218–223 fungi 58, 70, 91, 91, 100, 136, 139, 222, 278
birds 114–115 “the Great Dying” 223 mycorrhizae 104–105, 104, 105
clutch size 101, 114–115
turtles 253
Egler, Frank 173
Egyptians, ancient 82
Ehrlich, Paul 56, 59, 134, 250
electron microscopy 81, 85, 90
F G
electronic waste (e-waste) 331
facultative siblicide 115 Gaia hypothesis 197, 210, 214–217
elephants 22, 62, 64, 109, 139
elk 65, 110 Faraday, Michael 233 Galdikas, Birute 338
Elliott, Christopher 173 Farman, Joe 260–261 game theory 154, 155
Elser, James 43, 74, 75 feedback loops 136, 217, 224–225 garlic mustard 272–273, 273
Elton, Charles 50, 51, 100, 108, 109, 110–111, fermentation process 102, 103 Gaston, Kevin 185
112, 130, 132, 270, 272 fertilizers 327 Gause, Georgy 42–43, 52–53, 112, 190
empathy 125 runoff 151, 234, 239, 269 Gause’s Law 52–53, 112
endangered species 93, 95, 312 field manipulation experiments 63 gazelles 47, 73
endotherms 126, 222 fieldwork 43, 54–55, 116–117 genetically modified food 36, 36
energy fig trees 65 genetically modified organisms (GMOs) 36
biomass energy 305 fig wasps 58 genetics 19, 29, 154
energy flow through ecosystems 134, finches 27, 27, 110, 193 gene mapping 123
138–139 fireflies 89 gene selfishness 38–39
energy transfer 113, 136 fires 137, 171, 318, 318 gene therapy 19, 35
Enlightenment 20, 298 Fisher, Ronald 19, 29, 30, 114 genetic drift 81
environmentalism 294–331 fishing genetic engineering 35, 296
early history of 296–299 fish farming 269 genetic markers 123
environmental ethics 306–307 harmful practices 207, 238 heredity 18, 19, 20, 21, 26, 28, 32–33
Green Movement 297, 299, 308–309 moratoriums and quotas 267, 268 human genome 19, 34, 37, 123
halting climate change 316–321 overfishing 93, 150, 207, 229, 250, 266–269 see also DNA
Man and the Biosphere Programme (MAB) fishing cats 97 geothermal energy 91, 304
310–311 fixed action patterns (FAPs) 116–117 germ theory of disease 70, 102, 103
renewable energy 300–305 Flannery, Tim 339 Gessner, Conrad 80, 82, 83
Sustainable Biosphere Initiative (SBI) fleas 112 giant pandas 51, 51
322–325 Fleming, Alexander 102, 103 Gillespie, Rosemary 339
waste disposal 330–331 flight, insects 126–127 giraffes 18, 21
epidemics 71 flooding 238, 239, 258, 277 glaciation 198–199
epiphytes 169, 169 food chains 69, 75, 94, 108, 130, 132–133, 277 glaciers 198, 199, 199, 203
Esmark, Jens 198 DDT biomagnification 246 Gleason, Henry 152, 161, 171, 172, 174–175
essentialism 18, 20 ecological pyramids 112, 112 Global Biodiversity Information Facility (GBIF)
ethology 101, 116–125 producers and consumers 112, 132–133, 139, 180, 183
animal models and human behavior 101, 277, 279 global warming 185, 202–203, 207, 223, 224,
118–125 food limitation hypothesis 115 268–269, 276, 281, 318, 319
eukaryotes 90, 91 food webs 108, 108, 109, 133, 138, 140, 141, 142 goldfinches 181
eusocial species 39 Forbes, James 199 Gondwana 213, 222, 223
eutrophication 151 Forbes, Stephen A. 160, 166 Goodall, Jane 101, 120–122, 121, 122, 124, 125
INDEX 347
JK
Gore, Al 244, 304, 309, 319 HFCs 261
gorillas 94, 123 hibernation 278
Gosling, Raymond 19 Hoffman, Paul F. 337–338
Gould, Stephen Jay 38 Holdridge, Leslie 197, 206, 209
“great chain of being” 83 holistic theory 175, 210–211
Great Lakes, North America 150–151 Hölker, Franz 252 jaguars 65
Great Oxygenation Event 189 Holling, Crawford 131, 150–151 Janzen, Daniel 43, 55, 56–57
Great Pacific Ocean Garbage Patch (GPOGP) Holmes, Arthur 212 Janzen–Connell hypothesis 55
284 Holyoak, Marcel 190 Jenner, Edward 84, 102
Great Tits 279, 279 homeostasis 215, 217 Johnson, Roswell Hill 50
Green Movement 297, 299, 308–309 hominids 124 Johnston, Emma 235, 235
Green Revolution 327 Hooke, Robert 22, 42, 80, 84, 85, 85, 102 Jones, Clive 189
greenhouse gases and greenhouse effect 95,
hornets 127
153, 202, 202, 203, 228, 239, 240, 241, 264,
horses 31, 89 Kaminskyj, Susan 339
294, 299
Hubbell, Stephen P. 152, 190 kangaroo rats 185
Greenpeace 299, 309, 330
human behavior, animal models and 101, 118–125 Keeling, Charles 202, 228, 240
Grew, Nehemiah 85
human genome 19, 34, 37, 123 Keeling Curve 240–241, 241
Grimm, Nancy 338–339
Humboldt, Alexander von 42, 72, 160, 162–163, Kelly, Allan O. 220
Grinnell, Joseph 42, 50–51, 108, 110, 112, 176
163, 166, 168, 174, 176, 206, 256 Kelly, Petra 308, 309, 309
Grisebach, August 172
hummingbirds 110, 110 kelp 64–65, 143
Growth Rate Hypothesis (GRH) 75
Hunter, Tim 253 kestrels 111
guilds 152, 161, 176–177
hunting behavior 124 Keto, Aila 338
gulls 101, 117
Hutchinson, George Evelyn 50, 51, 52, keystone species 43, 60–65, 109, 130, 142
111, 139 kin selection 19, 29, 39
Klein, Naomi 262, 263, 263
H
Hutton, James 18, 23, 196, 198, 204
Huxley, Julian 19, 26, 86 Klopfer, Peter H. 336
hydroelectric power 294, 302, 304–305 koala bears 111, 111
Koch, Robert 100, 102
hydrosphere 197, 204, 215
Kolbert, Elizabeth 92, 202, 222
Krakatua 149, 149
habitats Krebs, Charles 224
I
carrying capacity 47 Kyoto Protocol 153, 320
destruction 93, 94, 95, 124, 137, 239, 280
endangered 236–239
L
fragmentation 93, 124, 130, 157
protected 239
Haeckel, Ernst 91, 166, 206, 334–335 ibis 97
Hairston, Nelson 130, 141 ice ages 198–199
Hamilton, William D. 19, 29, 38, 39, 154 ice cap melting 225, 318
Hansen, James 225 idealization of nature 298, 299 Lack, David 101, 114, 115
Hanski, Ilkka 161, 187, 187 immunity 70 Lack’s principle 115
Hardin, Garrett 108, 229, 250, 306 imperial ecology 296, 299 ladybugs 224, 271
Hardy, Sarah 339 lake ecosystem 211
imprinting 116, 117
hares 110, 110, 188 Laland, Kevin 188
inbreeding 314
Harkin, James Bernard 335 Lamarck, Jean-Baptiste 18, 20–21, 21, 26, 28, 32
Indo-Burma biodiversity hotspot 96, 97
Harrison, Nancy 189 land ethic 294–295, 306
industrial melanism 31, 31
Hartig, Theodor 104 landfill sites 331
Industrial Revolution 20, 31, 228, 232, 241,
Hatton, Harry 54 Lawton, John 189
294, 296
hawk-dove “game” 155 Leakey, Louis 120, 121
Ingersoll, Andrews 224
Hawking, Stephen 37 Leakey, Mary 336
insects
Hawkins, Charles 161, 177 Leeuwenhoek, Antonie van 42, 84–85, 100,
HCFCs 261 mass extinction 223 102, 130, 132
Heath Hen 313, 313 thermoregulation 126–127 Leibold, Mathew 190, 192, 193
heavy metals 105 interbreeding 88, 89 Lenski, Richard 31
Heinrich, Bernd 101, 126 intermediate disturbance hypothesis (IDH) 55 Leonardo da Vinci 22
Hennig, Willi 81, 90 International Dark-Sky Association 229, 253 Leopold, Aldo 140, 142, 167, 244, 294, 297, 306,
hens 165, 313 Internationl Union for Conservation of Nature 307, 307
herbivores 109, 113, 133, 139, 142 (IUCN) 236 Levin, Simon A. 338
heredity 18, 19, 20, 21, 26, 28, 32–33 invasive species 93, 148, 270–273 Levins, Richard 52, 186
Herodotus 42 animals 270–271 Lewontin, Richard 188
Hess, Harry 212 plants 272–273, 282 lichens 171
heterotherms 101, 126 island biogeography 94, 130–131, 144–149, life zone classification 197, 209
heterotrophs 133 193 ligers 89
348 INDEX
light pollution 235, 252–253 Mendel, Gregor 19, 26, 29, 32–33, 33, 296 nest predation hypothesis 114, 115
Likens, Gene 229, 248, 249, 249 Mendes, Chico 256, 257, 257, 258 Nestler, Johann Karl 33
Lindeman, Raymond 112, 113, 130, 138–139 metacommunities 190–193, 192 Neumann, John von 154
Linnaeus, Carl 20, 42, 80–81, 82, 83, 86–87, 87, 91, metamorphosis 77, 276, 279 Newport, George 101, 126
120, 132, 133, 162, 168 metapopulations 161, 186–187, 190 nitrogen 74
lions 49, 109 methane 331 noise pollution 235
Lithops (“flowering stones”) 168 Mexico City 283 noosphere 205
lithosphere 197, 204, 212 miasma 69 Nordhaus, William 324, 325, 325
lizards 112 mice 70, 71 nuclear power 217, 235, 302, 308
Locke, Harvey 339 microbes 90, 91, 102–103
locusts 75 microbial resistance 103
O
Loreau, Michel 131, 156–157 microhabitats 147
Lorenz, Edward 184 microplastics 284
Lorenz, Konrad 101, 116, 117, 117, 120, 123 microscopy 80, 84–85, 100, 102
Lotka, Alfred J. 42, 46, 47, 52, 224–225 Miescher, Friedrich 26, 32
Lotka–Volterra equations 42, 46–49, 52, 225 migration
Lovelock, James 197, 204, 210, 214, 215, 215, 216, birds 180, 180, 199, 278 oak trees 171, 189, 277
322 butterflies 181–182 obligate siblicide 115
Lowe, Ian 338 Miller, Brian 62, 65 oceans
Lubchenco, Jane 322, 323, 323 Miller, G. Tyler 137 acidification 207, 238, 281
Lyell, Charles 18, 23, 26, 196 Miller, Hugh 221 garbage patches 183, 284–285
lynx 48, 77, 110, 110, 188 Miner, Jack 335 octopus 83
“missing link” 121
Odling-Smee, John 161, 188, 189
Möbius, Karl August 334
Odum, Howard and Eugene 134, 138, 197, 210,
M
moles 21
210–211, 214
Molina, Mario 229, 260, 261
Odum, William E. 43
monarch butterflies 181–182, 182
Ohta, Tomoko 337
monoculture plantations 256
oil extraction 262, 263–264, 263
monsoons 291, 318
oil spills 234–235
MacArthur, Robert 43, 52, 53, 66, 131, 146–147, Montreal Protocol 260, 261, 319
one-child policy (China) 251
147, 150, 312 Moore, Charles J. 284, 285
open community theory 174–175
macaws 111 Morgernstern, Oskar 154
optimal foraging theory (OFT) 43, 66–67
McCallum, Malcolm 280 Morris, Desmond 116, 120, 122
orangutans 123
McClintock, Barbara 335 Morrone, J.J. 200
McKendrick, Anderson Gray 68, 164 orcas 234
mosquitoes 127, 247, 253
McKibben, Bill 264 orchids 59
mosses 169, 171
MacMahon, James 161, 177 Ortelius, Abraham 212
moths 31, 31, 56, 57, 59, 101, 126, 253, 273
macroecology 185 ospreys 247
mountain goats 191
Malle, Adolphe Dureau de la 170, 171 overgrazing 93, 140, 239, 265
mudslides 258, 258
Malthus, Thomas 18, 27, 46, 47, 164, 165, 165, overharvesting 93, 94–95
Muir, John 228, 236, 237, 237, 298, 306
184, 250 overpopulation 229, 250–251
mules 89
mangroves 146, 147, 239, 259 see also population dynamics
Munroe, Eugene 146
Marae Moana 239 musk oxen 72, 239 Owen, Richard 22
marginal value theorem (MVT) 66, 67 mussels 63, 67, 272 owls 111
Margulis, Lynn 204, 210, 215, 322, 337 mutualism 42, 43, 56–59, 100, 104, 105, 157 oxpeckers 58, 110
marine conservation 182–183, 239, 267 service-resource relationships 58 oystercatchers 66, 67, 67
marmots 278 service-service relationships 58, 59 ozone depletion 260–261, 319
Marsh, George Perkins 134, 135, 294, 299, 299 mycelium 104 ozone emissions 93
marsupials 209, 213 mycorrhizal fungi 104–105, 104, 105
mathematical modeling 54, 70, 74, 146–147, Myers, Norman 81, 96–97, 97
P
155, 184
matriarchal societies 125
N
Matthews, Blake 188
Mauna Loa 241, 241
Maupertuis, Pierre Louis Moreau de 20
Maurer, Brian 185, 338 Paczoski, Jósef 335
May, Robert 68, 70–71, 108, 150, 184 Paine, Robert 43, 54, 62–63, 63, 76, 130, 140,
mayflies 85 national parks 236, 237, 237, 239, 298, 307 141
Mayr, Ernst 81, 88 natural resources, depletion of 262–265 Pangaea 212–213, 223
meadowlarks 89 natural selection 18–19, 21, 22, 24–31, 38, 66, 72, parasites 49, 68, 71, 112, 187
meat-eating 109, 123–124, 133 81, 154 parasitoids 49, 49
megacities 282 Nelson, Gaylord 211, 295 Paris Agreement 318, 320–321, 321
megatsunami 221 nematodes 143 Parmesan, Camille 277, 277, 278
INDEX 349
S
parrotfish 63–64 population dynamics cont.
Pasteur, Louis 70, 100, 102, 103, 103 urbanization 282–283, 297
pasteurization 103 Verhulst equation 164–165, 184
peacocks 28, 29 population viability analysis (PVA) 312–315
peak oil 263–264 prairie dogs 62, 62, 63
Pearl, Raymond 164–165 Pratchett, Terry 120 Saint-Hilaire, Etienne Geoffroy 20
pedosphere 215 predators and prey 42, 56 salamanders 63, 283, 283, 291
Peltier, Autumn 339 apex predators 65, 76, 109, 133 salmon 184
penguins 72–73, 73 food chains 109 salt-tolerant vegetation 169
penicillin 102 nonconsumptive effects (NCEs) 76–77 saltmarshes 142, 210
Peregrine Falcons 229 predator–prey equations 44–49, 225 saolas 97
periphyton 113 trophic cascades 62, 130, 140–143 sawflies 278
permafrost 225 prickly pears 263 scala naturae 82, 83
pesticides 229, 242–247 primary consumers 112, 133, 138, 139, 277, Schimper, Andreas 160, 168, 169
phenology 276 279 Schmidt-Nielsen, Knut 72, 73, 73
phenotypes 30, 33, 89 primate development 120 Schumacher, Ernst 295, 328
photosynthesis 66, 72, 74, 133, 136, 189, 221,
prokaryotes 90 scientific revolution 296
222, 228, 240, 249, 259, 269, 303, 305
protists 91 Sclater, Philip 160, 162, 200
photovoltaic cells 302
ptarmigans 315 sclerophyll 169
phytogeography 200
Pulliam, Robert 66, 67 sea anemones 59, 59
phytoplankton 74, 94, 105, 112–113, 142, 222,
sea levels, rising 203, 225, 241, 318
269, 269, 281
QR
sea otters 64–65, 143
phytotrons 156, 157
sea snails 182
Pianka, Eric 66
sea urchins 64, 143
pioneer plants 160, 171
seals 109, 235
Pitton de Tournefort, Joseph 86
plant communities 160–161, 167 Sears, Paul 134
climax communities 172–173, 174 queen conches 182 seasonal cycle 276–279
open community theory 174–175 second-order predation hypothesis 143
plant distribution 168–169 secondary consumers 138, 139, 277
rabbits 48, 109, 132, 142, 225, 270–271
plant ecology 167 seeds
raccoons 111
plant physiology 169 dispersal 58, 64
rainforests 54, 55, 97, 153, 209, 209, 228, 238,
plant succession 135, 160, 167 diversity 326–327
256–257, 258, 259, 264
plants, invasive 272–273, 282 sessile species 77
rainwater catchment and storage 291
plastic pollution 232, 235, 269, 284–285 sewage treatment 228, 233, 291
ranaviruses 280
plate tectonics 212–213 sexual selection 28
rats 111
Playfair, John 23 Shachak, Moshe 189
Raven, Peter 56, 59
poaching 95 Shaffer, Mark 313, 314
Ray, John 80, 82, 83, 86, 88
polar bears 72, 109, 109 sharks 133
recycling 291, 330–331
politics, Green 308, 309 Shaw, Robert 337
Red List of the International Union for
pollination 56, 57, 58, 59, 230–235, 250, 279, Shelford, Victor 138, 206, 207, 208
Conservation of Nature (IUCN) 93, 95, 312 Shiva, Vandana 326, 327, 327
329
Red Queen evolutionary theory 49 sickle cell disease 37, 37
pollution 93–94, 105, 225, 228, 229, 280
Redfield Ratio 74 Siemens, Werner von 302
acid rain 93, 222, 229, 234, 248–249
Reed, Lowell 164 Sierra Club 298, 307, 308
air 93, 95, 232, 233–234, 233, 248
reforestation 259 silver maple 209
effects on health 232, 234
Renaissance 296 Simpson, George Gaylord 212, 213
intangible pollutants 235
light 235, 252–253 renewable energy 300–305 Slagsvold, Tore 114, 115
noise 235 resource partitioning 53 Slobodkin, Lawrence 130, 140, 141, 143
oil spills 234–235 rhinos 201, 223 Smith, Frederick 141
pesticides 229, 242–247 ribosomes 90–91 Smith, John Maynard 29, 48, 131, 154–155
plastic 232, 235, 269, 284–285, 284, 285 rice 36 Smith, Robert Angus 248
thermal 235 Richmond, William Blake 335 smog 233
water 93, 94, 228, 233, 234–235, 269, 289, 330 Rio Earth Summit 295 snakes 93
population dynamics 46–49, 108, 110 RNA 36 Snow, John 69–70, 69
chaotic change 184 ribosomal RNA (rRNA) 75 snowshoe hares 110, 110
ecological drift 192 Robins 114 soil acidification 93
human population growth 27, 46, 47, 94, 164, rockpools 193 solar power 294, 302–303, 305
237 Romanticism 298, 299 Sonora Desert 173, 312
metapopulations 161, 186–187 Roosevelt, Theodore 264, 335 sparrows 253
overpopulation 229, 250–251 Root, Richard 152, 161, 176–177 speciation 30, 88–89
population control 250, 251 Rowland, Frank 229, 260, 261 species distribution 162–163
predator–prey equations 44–49, 190, 225 “runaway greenhouse” effect 224, 225 species-area effect 147
350 INDEX
Spencer, Herbert 172 trophic-dynamic theory 138–139 wasps 49, 58

spontaneous generation 102 trout 111, 211 waste disposal 330–331
spring creep 274–279 truffles 104 water
squirrels 53, 53, 278 turtles 182, 229, 239, 253, 253, 285 economic water scarcity 290–291
starfish 62, 63, 130, 141, 141 Tyndall, John 202, 203 hydroelectric power 294, 302, 304–305
starlings 66, 67, 111, 189 typological species concepts 89 physical water scarcity 290
Steller’s sea cows 143, 143 pollution 93, 94, 228, 233, 234–235, 269, 289, 330
U
steppes 207 recycling 291
Sterner, Robert 43, 74 saline 265, 288
sticklebacks 117 tidal power 302, 305
stinkhorn fungi 58 water crisis 286–291
stochasticity 192, 313, 314 water stress 290
stromatolites 205 water supplies 265, 288, 289–290
Udvardy, Miklos 200, 209
Suess, Eduard 134, 197, 204 Watson, Andrew 216
UNESCO 310, 311
sugar maple 209 Watson, James 19, 32, 34–35, 35
uniformitarianism 18, 23
Sugihari, George 184 weasels 109
urbanization 282–283, 297
survival of the fittest 28, 37, 53 Wecker, Stanley C. 337
Urqhuart, Fred and Norah 180, 181, 181, 182
Suzuki, David 337 Wegener, Alfred 163, 196, 212–213
UV radiation 260, 261
swallows 199 Werner, Abraham 23
Swammerdam, Jan 85 Werner, Earl 43, 76–77
V
symbiotic relationships 56 wetlands 239, 289, 291
systems ecology 211 whales 87, 133, 235, 239, 285
whelks 62, 63
White, Gilbert 294, 297, 298
T
White, Lynn 306
vaccines 84, 102–103 Whitehouse, Michael 189
Van Valen, Leigh 46, 49 Whittaker, Robert 86, 90, 91, 161, 174, 175, 209
vegetation Wickett, Michael E. 281
biomes 208–209 Wiesenfeld, Kurt 184
tadpoles 76–77, 109 climax communities 172–173 wildlife corridors 239
taiga 201 feedback loops 224 wildlife crossings 191, 191
Tang, Chao 184 formations 173, 206–207 Wilkins, John 85
Tansley, Arthur 130, 134, 135, 135, 136, 138, 153, open community theory 174–175 Wilkins, Maurice 35
167, 172, 174, 176, 190, 208, 210, 214 plant distribution 168–169 Williams, Carrington 185
tapirs 163 plant succession 135, 160, 167 Williams, George C. 38
tarns 136 zones 168–169 Wilmut, Ian 34
taxonomy 37, 80–81, 86–87 see also deforestation; plant communities Wilson, E.O. 81, 92, 93, 94, 94, 131, 146, 147, 148,
Teale, Edwin Way 246 Verhulst, Pierre-François 164, 184 149, 223, 312
telomeres 123 Verhulst equation 164–165, 184 wind power 303–304, 323
Ten Percent Law 113 Vernadsky, Vladimir 136, 138, 153, 160, 167, 197, Wingfield, Eileen Wani 336
tench 113 204–205, 205, 214 Woese, Carl 81, 90–91
territorial behavior 155 viral shunt 69 wolves 49, 64, 65, 110, 140
Theophrastus 42 viruses 68, 69, 70, 102, 103, 280 “wood-wide web” 105
thermal pollution 235 volcanic activity 149, 149, 163, 222, 223 woodpeckers 111, 189
thermoregulation 126–127 Volterra, Vito 46, 46, 47, 48, 52, 225 World Wide Fund for Nature (WWF) 236
Thoreau, Henry David 171, 244, 298 Wright, Judith 336
ticks 58, 112
W
tidal power 302, 305
XYZ
tigers 49, 133, 201
Tinbergen, Niko 39, 101, 116, 117, 123
toads 280
tool use, animal 120, 121, 122
topsoil erosion 264–265 Waddington, Conrad 188
tortoises 27, 191, 200 Wagler, Ron 223 X-ray crystallography 19
Traill, Catherine Parr 334 Wahlenburg, Göran 198
transition engineering 265 Wallace, Arthur Russel 18, 27–8, 29, 42, 59, 146, yellow bush lupines 142, 143
transmutation 18, 21, 28 160, 162, 163, 166, 196–197, 200–201, 201 Yorke, James A. 338
tree diversity 55 Wallace Line 163, 201 yuccas 57
Triassic Period 220, 223 Walter, Heinrich 206
trilobites 223 warblers 52, 53, 199 zoogeography 162, 163, 200, 200, 201
trophic cascades 62, 130, 140–143 warfare 124–125
trophic levels 130, 139, 141 “warm-blooded” 126
trophic pyramid 113 Warming, Johannes 160, 166, 167
351
QUOTE ATTRIBUTIONS
THE STORY ECOSYSTEMS THE HUMAN FACTOR
OF EVOLUTION
132 Richard Bradley 230 Barry Commoner
134 Tyler Miller and Scott Spoolman 236 John Muir
20 Jean-Baptiste Lamarck
152 Stephen Hubbell 240 Ralph Keeling
22 Georges Cuvier
153 Long Term Ecological Research (LTER) 242 Rachel Carson
23 James Hutton
Network 248 Gene Likens
24 Charles Darwin
156 Michel Loreau 250 Garrett Hardin
32 Haruki Murakami
252 Tim Hunter
34 Francis Crick
254 Chico Mendes
260 Carl Sagan
ORGANISMS 262 Gro Harlem Brundtland
ECOLOGICAL IN A CHANGING 266 Magraret Atwood
270 Thomas Austin
PROCESSES ENVIRONMENT 274 Jonathan Banks
280 Stephen Price
44 Vito Volterra 162 Alexander Von Humboldt 281 Elizabeth Kolbert
50 Joseph Grinnell 164 Pierre François Verhulst 286 Maude Barlow
52 Georgy Gause 166 Stephen Alfred Forbes
54 Joseph Connell 167 David Attenborough
56 Daniel Janzen
60 Kevin D. Lafferty and Thomas Suchanek
170 Henry David Thoreau
172 Frederic E. Clements
ENVIRONMENTALISM
66 Eric Charnov, H.R. Pulliam, and Graham Pyke 174 Henry Allan Gleason AND CONSERVATION
76 Liana Zanette 176 R.B. Root
178 Brent Mitchell
296 Francis Bacon
184 Lev R. Ginzburg
297 Gilbert White
185 James Brown
ORDERING THE 186 Richard Levins
298 Henry David Thoreau
299 George Perkins Marsh
NATURAL WORLD 188 John Odling-Smee, Kevin Laland, and Marcus
300 Werner von Siemens
Feldman
306 Aldo Leopold
82 Aristotle 310 UNESCO
84 Robert Hooke 312 Mark L Shaffer
86 Carl Linnaeus THE LIVING EARTH 316 Barack Obama
88 Ernst Mayr 324 William Nordhaus
90 George Fox and Carl Woese 326 Vandana Shiva
198 Louis Agassiz 328 Gretchen Daily
92 E.O. Wilson
202 James Hansen 330 Paul Connett
96 Norman Myers
204 Vladimir Vernadsky
210 Eugene Odum
212 Seth Shostak
THE VARIETY OF LIFE 214 James Lovelock
218 Walter Alvarez and Frank Asaro
224 James Hansen
102 Louis Pasteur
104 A.B. Frank
106 Charles Elton
116 Konrad Lorenz
118 Louis Leakey
352
ACKNOWLEDGMENTS
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РЕЛИЗ ПОДГОТОВИЛА ГРУППА "What's News" VK.

GENES ARE SOLAR ENERGY IS BOTH

THE TIME HAS

THE FOOD IS THE

DK US/LONDON ASSISTANT ART EDITORS

JACKETS EDITORIAL COORDINATOR Copyright © 2019 Dorling Kindersley Limited

EDI TORIAL DIRECTOR Printed and bound in Malaysia

DK DELHI Ellen Dupont A WORLD OF IDEAS:

CELIA COYNE TOM JACKSON

54 Poor field experiments 76 Fear itself is powerful

56 More nectar means

84 By the help of 138 Life is supported by a vast

88 “Reproductively isolated” 150 It is the constancy of

THE VARIETY depends on temperature

156 Species maintain the 170 I have great faith

A CHANGING 174 An association

190 Local communities that 214 Life changes Earth

218 65 million years ago 270 The introduction

FACTOR has been hurled against

260 The hole in the ozone

262 We needed a mandate

266 Bigger and bigger boats

308 Think globally, 326 Monocultures and

312 Predicting a population’s 330 We are living on this

299 Man everywhere is

300 Solar energy is

HMS Beagle sets sail on a

1785 1813 1831

Jean-Baptiste Lamarck Amateur fossil

THE STORY OF EVOLUTION 19

Gregor Mendel’s paper The Selfish Gene by

1859 1953 2003

Darwin elaborates on his In The Eagle pub in The Human Genome

THE STORY OF EVOLUTION 21

History in Paris, he argued that

1802 Research on the

THE STORY OF EVOLUTION 23

THE STRUGGLE FOR

26 EVOLUTION BY NATURAL SELECTION

Charles Darwin Born in Shropshire, UK, in 1809, instantly. Despite continuing

THE STORY OF EVOLUTION 27

of 1835 provided especially strong Comparison of Galapagos finch

28 EVOLUTION BY NATURAL SELECTION

There is variation There is differential reproduction. There is heredity. End result:

THE STORY OF EVOLUTION 29

Building on the theory

30 EVOLUTION BY NATURAL SELECTION

THE STORY OF EVOLUTION 31

THE STORY OF EVOLUTION 33

dominant or recessive. When both

Pioneering geneticist Gregor Johann Mendel

THE STORY OF EVOLUTION 35

College, London, Franklin and

Genetic engineering useful—have greatly simplified

36 THE ROLE OF DNA

Good and bad errors

THE STORY OF EVOLUTION 37

2002 Stephen Jay Gould

THE STORY OF EVOLUTION 39

circumstances in which the gene

Robert MacArthur’s Dan Janzen observes the

1917 1957 1965

1925–26 1961 1969