The Ecology Book PDF
The Ecology Book PDF
The Ecology Book PDF
COM/WSNWS
ECOLOGY
QUESTION
ALL BODILY
ACTIVITY DEPENDS
ON TEMPERATURE
BOOK
BIG IDEAS SIMPLY EXPLAINED
THINK GLOBALLY,
ACT LOCALLY WE ARE
LIVING
ON THIS
PLANET AS
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.
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.
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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
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
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
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INTRODU
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CTION
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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
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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,
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THE STO
OF EVOL
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RY
UTION
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18 INTRODUCTION
1809 1823
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.
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1866 1976
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. ■
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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
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Key works
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
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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
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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
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 ❯❯
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Natural selection
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
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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
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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
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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.
ECOLOGI
PROCESS
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CAL
ES
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42 INTRODUCTION
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
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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.
1972 1991
LESSONS FROM
MATHEMATICAL THEORY
ON THE STRUGGLE
FOR LIFE
PREDATOR–PREY EQUATIONS
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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
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ECOLOGICAL PROCESSES 47
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
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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
ECOLOGICAL PROCESSES 49
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. ■
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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.
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ECOLOGICAL PROCESSES 51
See also: Competitive exclusion principle 52–53 ■ Field experiments 54–55 ■ Optimal foraging theory 66–67
■ Animal ecology 106–113 ■ Niche construction 188–189
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. ■
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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.
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ECOLOGICAL PROCESSES 53
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
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.
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ECOLOGICAL PROCESSES 55
See also: Ecological niches 50–51 ■ Modern view of diversity 90–91 ■ Animal behavior 116–117 ■ The ecosystem 134–137
■ Niche construction 188–189
56
IN CONTEXT
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.
ECOLOGICAL PROCESSES 57
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 ❯❯
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58 MUTUALISMS
ECOLOGICAL PROCESSES 59
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. ■
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WHELKS
ARE LIKE LITTLE
WOLVES
IN SLOW MOTION
KEYSTONE SPECIES
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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.
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ECOLOGICAL PROCESSES 63
See also: Predator–prey equations 44–49 ■ Mutualisms 56–59 ■ Animal ecology
106–113 ■ Trophic cascades 140–143 ■ Evolutionarily stable state 154–155
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 ❯❯
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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
ECOLOGICAL PROCESSES 65
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-
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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,
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ECOLOGICAL PROCESSES 67
See also: Evolution by natural selection 24–31 ■ Predator–prey equations
44–49 ■ Competitive exclusion principle 52–53 ■ Mutualisms 56–59
68
IN CONTEXT
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
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ECOLOGICAL PROCESSES 69
See also: The microbiological environment 84–85 ■ Microbiology 102–103 ■ The ubiquity of mycorrhizae 104–105
■ Biodiversity and ecosystem functioning 156–157
10–15 deaths
Broad Street
pump
eet
d Str
B r oa
Re
ge
nt
S
t re
Fatalities in London’s et
et
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.
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ECOLOGICAL PROCESSES 71
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.
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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
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ECOLOGICAL PROCESSES 73
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
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
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ECOLOGICAL PROCESSES 75
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.
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.
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ECOLOGICAL PROCESSES 77
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
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
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ORDERIN
NATURAL
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G THE
WORLD
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80 INTRODUCTION
1665 CE 1758
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,
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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
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. ■
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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.
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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. ■
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.
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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
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86
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.
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Key works
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
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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.
92
IN CONTEXT
SAVE THE
KEY FIGURE
Edward O. Wilson (1929–)
BIOSPHERE
BEFORE
1993 The UN proclaims
December 29 as the
International Day for
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
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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.
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THE VARI
OF LIFE
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ETY
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100 INTRODUCTION
1676 1885
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.
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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. ■
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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.
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104
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
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FOOD IS THE
BURNING
QUESTION
ANIMAL ECOLOGY
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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
Dolphin
Copepod
Squid
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. ■
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114
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.
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116
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
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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
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REDEFINE “TOOL,”
REDEFINE “MAN,” OR ACCEPT
CHIMPANZEES
AS HUMANS
USING ANIMAL MODELS TO UNDERSTAND
HUMAN BEHAVIOR
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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.
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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 ❯❯
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
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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.
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126
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
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ECOSYST
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EMS
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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.
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
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ECOSYSTEMS 131
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. ■
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132
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
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”. ■
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134
IN CONTEXT
ARE POTENTIAL
(1871–1955)
BEFORE
1864 George Perkins Marsh,
ORGANISMS
Eduard Suess proposes
the term “biosphere.”
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.
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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
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 ❯❯
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.
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.
1957 American ecologist
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.
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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.
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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.
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
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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
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
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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
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ISLANDS ARE
ECOLOGICAL
SYSTEMS
ISLAND BIOGEOGRAPHY
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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
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ECOSYSTEMS 147
See also: Evolution by natural selection 24–31 ■ Predator-prey equations 44–49
■ Field experiments 54–55 ■ The ecosystem 134–137
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, ❯❯
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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
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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
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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. ■
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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
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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
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154
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
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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
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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
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ECOSYSTEMS 157
See also: Mutualisms 56–59 ■ Keystone species 60–65 ■ The ecosystem
134–137 ■ Organisms and their environment 166 ■ Invasive species 270–273
ORGANIS
IN ACHAN
ENVIRON
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MS
GING
MENT
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160 INTRODUCTION
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.
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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. ■
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162
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
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164
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
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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
166
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.
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
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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
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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
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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
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
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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
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174
AN ASSOCIATION
IS NOT AN ORGANISM
BUT A COINCIDENCE
OPEN COMMUNITY THEORY
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
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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.
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THE CITIZEN
NETWORK DEPENDS ON
VOLUNTEERS
CITIZEN SCIENCE
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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,
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Key works
Science should be
dominated by amateurship
instead of money-biased
technical bureaucrats.
Erwin Chargaff
Austro-Hungarian biochemist
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
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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
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186
A POPULATION
OF POPULATIONS
METAPOPULATIONS
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
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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
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188
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.
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190
IN CONTEXT
THAT EXCHANGE
BEFORE
1917 Arthur Tansley observes
that two species of Galium
COLONISTS
plants grow differently in
different soil patches.
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
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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.
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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
frog
heron cormorant
rushes heron frog
lake 3
perch mosquito
rushes algae carp cormorant
lake 4 lake 5
mosquito
duck algae
mayfly
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. ■
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THE LIVI
EARTH
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NG
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196 INTRODUCTION
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
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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.
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. ■
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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.
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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
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.
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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
n
r ad i a re d
E
atio
a Catastrophe, journalist
O
M
EARTH
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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,
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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
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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
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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.
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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.
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.
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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
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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.
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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
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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
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65 MILLION
YEARS AGO SOMETHING
KILLED HALF
OF ALL THE LIFE ON
THE EARTH
MASS EXTINCTIONS
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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
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
224
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
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THE HUM
FACTOR
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AN
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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.
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
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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.
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. ■
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ENVIRONMENTAL
POLLUTION IS AN
INCURABLE
DISEASE
POLLUTION
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232 POLLUTION
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
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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
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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. ■
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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
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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 ❯❯
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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
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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.
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THE CHEMICAL
BARRAGE HAS
BEEN HURLED AGAINST THE
FABRIC OF LIFE
THE LEGACY OF PESTICIDES
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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
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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.
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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. ■
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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.
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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
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I AM
FIGHTING FOR
HUMANITY
DEFORESTATION
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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
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
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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,
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260
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
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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. ■
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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
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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 ❯❯
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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
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266
IN CONTEXT
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
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Bycatch
Physical impact
of fishing gear
270
IN CONTEXT
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.
AS TEMPERATURES INCREASE,
THE DELICATELY
BALANCED SYSTEM
FALLS INTO
DISARRAY
SPRING CREEP
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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.
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
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Decoupling
Some organisms’ survival could be
threatened by the “decoupling” of
interactions between species. This
280
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
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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.
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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.
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284
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.
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WATER
IS A PUBLIC
TRUST, AND A
HUMAN RIGHT
THE WATER CRISIS
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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
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97.5% 79%
Oceans Ice caps and
glaciers
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
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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
KEY
None
1–9 months
ENVIRONM
AND CONS
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ENTALISM
ERVATION
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294 INTRODUCTION
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.
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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. ■
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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
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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. ■
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.
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
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SOLAR
ENERGY
IS BOTH WITHOUT LIMIT
AND WITHOUT
RENEWABLE ENERGY
COST
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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.
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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.
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306
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
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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. ■
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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
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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
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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.
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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.
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 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,
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A population A management
viability analysis is solution is found to
conducted to assess combat the threat
the situation. to the population.
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. ■
CLIMATE
CHANGE
IS HAPPENING HERE.
IT IS HAPPENING
HALTING CLIMATE CHANGE
NOW
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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
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. ■
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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
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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.
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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.
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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
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330
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
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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
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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
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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
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
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
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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.
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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.
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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.
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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.
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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
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
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INDEX 345
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
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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
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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
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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
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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
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350 INDEX
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
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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
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352
ACKNOWLEDGMENTS
Dorling Kindersley would like to thank Professor Fred Terry Whittaker Wildlife (br). 127 Alamy Stock Photo: (bc). 239 Alamy Stock Photo: ImageBroker (crb); Huang
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