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Shreya Singal, Vidushi Gupta, Amrai Dua
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Julia Schroeder received her Ph.D. in Animal Ecology from the A naturalist and teacher with a particular interest in evolutionary
University of Groningen in the Netherlands. From 2012 to 2017, biology, Derek Harvey graduated in Zoology from Liverpool University
she headed a research group at the Max Planck Institute for in the UK. He has taught a generation of biologists and led student
Ornithology in Germany, studying social behavioral ecology. Julia expeditions to Costa Rica, Madagascar, and Australasia. Derek now
currently researches and teaches evolutionary biology at Imperial concentrates on writing and consulting for science and natural
College London. history books.


Celia Coyne is a freelance writer and editor living in Christchurch, A writer for 25 years, Tom Jackson is the author of about 200 nonfiction
New Zealand. She is the author of Earth’s Riches and The Power of books for adults and children and has contributed to many more. Tom
Plants and writes and edits articles on science and natural history studied zoology at Bristol University, UK, and worked in zoos and as a
for magazines, newspapers, journals, websites, and books in the UK, conservationist before turning to writing about natural history and all
Australia, and New Zealand. Her aim is to make scientific subjects things scientific.
accessible to lay readers.
Alison Singer is a Ph.D. candidate in Community Sustainability at
The author of hundreds of books on science and nature for both Michigan State University, US, where she studies storytelling and
children and adults, John Farndon studied geography at Cambridge science communication. She has a broad educational background in
University. He has written extensively on earth sciences and the writing, ecology, and the social sciences. Alison has worked as an
environment, focusing in particular on conservation and ecology. educator for environmental charities, and for the US Environmental
His books include The Oceans Atlas, The Wildlife Atlas, How the Earth Protection Agency.
Works, and The Practical Encyclopedia of Rocks and Minerals.

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.

12 INTRODUCTION 34 We’ve discovered
the secret of life
The role of DNA
OF EVOLUTION 38 Genes are selfish
The selfish gene
20 Time is insignificant
and never a difficulty
for nature
Early theories of evolution ECOLOGICAL
22 A world previous to ours,
destroyed by catastrophe 66 The fitness of a foraging
Extinction and change 44 Lessons from animal depends on
mathematical theory its efficiency
23 No vestige of a beginning on the struggle Optimal foraging theory
—no prospect of an end for life
Uniformitarianism Predator–prey equations 68 Parasites and pathogens
control populations
24 The struggle for existence 50 Existence is determined like predators
Evolution by natural selection by a slender thread Ecological epidemiology
of circumstances
32 Human beings are Ecological niches 72 Why don’t penguins’
ultimately nothing feet freeze?
but carriers for genes 52 Complete competitors Ecophysiology
The rules of heredity cannot coexist
Competitive exclusion 74 All life is chemical
principle Ecological stoichiometry

54 Poor field experiments 76 Fear itself is powerful

can be worse than Nonconsumptive effects
useless of predators on their prey
Field experiments

56 More nectar means

more ants and more ORDERING THE
ants mean more nectar
60 Whelks are like 82 In all things of nature
little wolves in there is something of
slow motion the marvelous
Keystone species Classification of living things

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

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

88 “Reproductively isolated” 150 It is the constancy of

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

THE VARIETY depends on temperature

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

156 Species maintain the 170 I have great faith

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

A CHANGING 174 An association

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

190 Local communities that 214 Life changes Earth

exchange colonists to its own purposes
Metacommunities The Gaia hypothesis

218 65 million years ago 270 The introduction

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

FACTOR has been hurled against

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

260 The hole in the ozone

layer is a kind of
Ozone depletion

262 We needed a mandate

for change
Depletion of natural resources

266 Bigger and bigger boats

chasing smaller and
fewer fish

306 The time has come for 324 We are playing dice with
science to busy itself the natural environment
with the Earth itself The economic impact of
Environmental ethics climate change

308 Think globally, 326 Monocultures and

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

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

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

299 Man everywhere is

a disturbing agent
Human devastation
of Earth

300 Solar energy is

both without limit
and without cost
Renewable energy

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




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


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,




HMS Beagle sets sail on a

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

1785 1813 1831

1809 1823

Jean-Baptiste Lamarck Amateur fossil

publishes Philosophie Zoologique, hunter Mary Anning
where he argues that animals acquire uncovers the first intact
characteristics as a consequence of plesiosaurus skeleton.
use or nonuse of different body parts,
triggering mutations over generations.

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.


Gregor Mendel’s paper The Selfish Gene by

“Experiments with Plant Hybrids” evolutionary biologist Richard
outlines findings from his pea Dawkins offers a new
plant experiments, laying perspective on evolution,
the foundations for the looking at the gene, as opposed
field of genetics. to the species or group.

1866 1976

1859 1953 2003

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

theories of evolution in On the Cambridge, UK, Crick and Project produces the first
Origin of Species by Means Watson announce that genetic blueprint of
of Natural Selection, which they have discovered Homo sapiens.
is an instant sellout. the structure of DNA.

Although the idea of evolution of thought were complementary, that genetic information is “written”
became widely accepted, the rather than contradictory. In 1942, on DNA molecules. The errors that
mechanism that made natural Julian Huxley articulated the occur when DNA copies itself create
selection possible was not yet synthesis between Mendel’s mutations—the raw materials for
known. In 1866, an Austrian monk genetics and Darwin’s theory evolution. By the 1980s it was
called Gregor Mendel made a huge of natural selection in his book possible to map and manipulate the
contribution to genetics when he Evolution: The Modern Synthesis. genes of individuals and species. In
published his findings on heredity the 1990s, the mapping the human
in pea plants. Mendel described The double helix genome paved the way for medical
how dominant and recessive traits Advances in technology such as research into gene therapy.
pass from one generation to the X-ray crystallography led to more Ecologists also want to establish
next, by means of invisible “factors” discoveries in the 1940s and ’50s, whether genes influence behavior.
that we now call genes. and the foundation of the new Back in 1964, William D. Hamilton
The rediscovery of Mendel’s discipline of molecular biology. popularized the concept of genetic
work in 1900 initially sparked sharp In 1944, chemist Oswald Avery relatedness (“kin selection”) to
debate between his supporters and identified deoxyribonucleic acid explain altruistic behavior in
many Darwinians. At the time, (DNA) as the agent for heredity. animals. In The Selfish Gene (1976),
evolution was believed to be based Rosalind Franklin and Raymond Richard Dawkins further advanced
on the selection of small, blending Gosling photographed strands of the the gene-centered approach. It is
variations, but Mendel’s variations DNA molecule in 1952, and James clear that aspects of evolutionary
clearly did not blend. Three decades Watson and Francis Crick confirmed biology will still spark debate as
later, geneticist Ronald Fisher and its double helix structure the long as ecologists continue to
others argued that the two schools following year. Crick then showed develop Darwin’s theory. ■



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
unchanged throughout time—a view fact). As Earth cooled, species had
The Comte de Buffon
now known as essentialism. This appeared, died off, and were finally
(1707–88), Jean-Baptiste
idea came under challenge as a replaced by ancestors of those
Lamarck (1744–1829) result of two developments: the known today. Noting similarities
BEFORE intellectual movement known as the among animals such as lions,
1735 Swedish botanist Carl Enlightenment (c. 1715–1800), and tigers, and cats, Buffon deduced
Linnaeus publishes Systema the Industrial Revolution (1760–1840). that 200 species of quadrupeds had
Naturae, a system of biological The Enlightenment was marked evolved from just 38 ancestors. He
classification that later helped by scientific progress and increased also believed that changes in body
to determine species’ ancestry. questioning of religious orthodoxy, shape and size in related species
such as the claim that God created had occurred in response to living
1751 In “Système de la nature” Earth and all living things in seven in different environments.
French philosopher Pierre days. Then, as the Industrial In 1800, French naturalist Jean-
Louis Moreau de Maupertuis Revolution gathered pace, canals, Baptiste Lamarck went further. In a
introduces the idea that railroads, mines, and quarries lecture at the Museum of Natural
features can be inherited. cut through rock strata and revealed
thousands of fossils, mostly of
AFTER animal and plant species that no
1831 Etienne Geoffroy Saint- longer existed and had never been
Hilaire writes that sudden seen before. These suggested that
environmental change can life began long before the widely
cause a new species to develop accepted creation date of 4400 bce, Nature is the system of laws
from an existing organism. deduced from biblical sources. established by the Creator for
the existence of things and
1844 In Vestiges of the Natural Animal adaptation for the succession of creatures.
History of Creation, Scottish In the late 1700s, French scientist The Comte de Buffon
geologist Robert Chambers Georges-Louis Leclerc, Comte de
argues—anonymously—that Buffon, upset church authorities
simple creatures have evolved by asserting that Earth was much
into more complex species. older than the Bible suggested. He
believed it was formed from molten


See also: Extinction and change 22 ■ Uniformitarianism 23 ■ Evolution by
natural selection 24–31 ■ The rules of heredity 32–33

History in Paris, he argued that

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

Key works

1802 Research on the

Organization of Living Bodies
1809 Zoological Philosophy
1815–22 Natural History of
Invertebrate Animals



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
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



niformitarianism is the
IN CONTEXT theory that geological
processes, such as the
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




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

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

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


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

of 1835 provided especially strong Comparison of Galapagos finch

evidence for his later theory of bill structure
evolution by natural selection. Here,
he observed that the shape of the
carapaces (shells) of giant tortoises
varied slightly from island to island.
Darwin was also intrigued to find
that there were four broadly similar,
yet clearly distinct, varieties of
mockingbirds, but that no single
island had more than one species Geospiza magnirostris Geospiza fortis
The short, sharp bill of the Large The bill of the Medium Ground Finch
of the bird. He saw small birds, Ground Finch, the biggest of Darwin’s is variable, evolving rapidly to adapt
too, that looked alike but had a finches, enables it to crack nuts. to whatever size seeds are available.
range of beak sizes and shapes.
Darwin deduced that each group
possessed a common ancestor but
had developed diverse traits in
different environments.

Darwin’s conclusions
On Darwin’s return to England, the Geospiza parvula Certhidea olivacea
differing beaks of the small birds The stubby bill of the Small Tree Finch, The slender, probing bill of the Green
which forages in foliage, suits its diet Warbler-finch helps it catch small
he had found on the Galapagos, of seeds, fruits, and insects. insects and spiders.
usually called “finches” although
they are not in the true finch family,
set him thinking. He knew that populations had evolved in different Malthus predicted that population
a bird’s beak is its key tool for Galapagos habitats, each group growth would eventually outstrip
feeding, so its length and shape adapted for a more or less specialist food production. This idea matched
offer clues to its diet. Later research diet by a process that he would the evidence Darwin had observed
revealed that there are 14 different later call “natural selection.” Over of ongoing competition between
finch species on the Galapagos time, the finch populations had individual animals and species for
islands. The differences in their become distinct species. resources. This competitive aspect
beaks are marked and significant. In the early 21st century, formed the backbone of Darwin’s
For example, cactus finches have researchers at Harvard University coalescing theory of evolution.
long, pointed beaks that are ideal uncovered new evidence of how By 1839, Darwin had developed
for picking seeds out of cactus this happens at a genetic level. an idea of evolution by natural
fruits, while ground finches have Their findings, published in 2006, selection. He was, though, reluctant
shorter, stouter beaks that are showed that a molecule called to publish because he understood
better suited for eating large seeds calmodulin regulates the genes that the theory would unleash a
on the ground. Warbler finches have involved in shaping birds’ beaks, storm of controversy from those
slender, sharp beaks, which are and is found at higher levels in who would view it as an attack
ideal for catching flying insects. longer-beaked cactus finches than on religion and the Church. When,
Darwin speculated that the in shorter-beaked ground finches. in 1857, he began receiving
finches were descended from a communications from fellow British
common ancestral finch that had Refining the theory naturalist Alfred Russel Wallace,
reached the archipelago from the Darwin was influenced by Thomas who had independently arrived at
mainland of South America. He Malthus’s An Essay on the Principle very similar conclusions, Darwin
concluded that a variety of finch of Population (1798), in which realized he had to publish his ❯❯


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

Natural selection

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

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


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

Building on the theory

Many of Darwin’s and Wallace’s
ideas have proved remarkably
accurate, despite the fact that Why do some die and some
the workings of genetics were not live?… the answer was
understood at the time. Although clearly, that on the whole the
Darwin himself had used the best fitted live.
term “genetic” as an adjective Alfred Russel Wallace
to describe the as-yet-unknown In honeybee colonies, female
mechanism of inheritance, it was worker bees look after the queen
bee. They build the honeycomb,
British biologist William Bateson, gather nectar and pollen, and feed
in the early 20th century, who first larvae, but they do not breed.
used the term “genetics” in a


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


Evolution in real time
Richard Lenski, a professor
Individuals within at Michigan State University,
a species have established the Long-term
a variety of forms Experimental Evolution
Seen in the light of evolution, of a characteristic. project in 1988. For more than
biology is, perhaps, 25 years, he studied 59,000
intellectually the generations of the E. coli
most satisfying and bacterium. During this time,
inspiring science. he observed that the species
Theodosius Dobzhansky used the glucose solution
it lived in more efficiently,
increasing in size but also
The individuals with
growing faster. Also, a new
the characteristic best
species had evolved that was
suited to the environment
able to use a compound in the
are more likely to survive
solution called citrate, which
and breed.
the parent bacterium could
occurred. For example, a fossil
not. Evolving bacteria can
record stretches back 60 million
pose a potential threat to
years for ancestors of the horse. humans. Increasing antibiotic
The earliest of these were dog- use destroys many disease-
sized forest-dwelling animals with causing bacteria, but not those
several toes on each foot. Evolution with mutations that make
produced much larger horses with These them resistant to the drugs.
just a single hoof on each foot, characteristics As the non-resistant bacteria
adapted for life on open grasslands are passed on are killed off, the resistant
where they would often have had to the next strains become more
to outrun predators. generation. dominant, multiplying and
Peppered moths (biston passing on their mutations
betularia) reveal change over a to future generations. That
shorter period. The moth is usually is natural selection at work.
pale, providing camouflage against
the bark of birch trees, but a
mutation produces some black
moths. Before the 19th century,
most peppered moths were pale.
During the Industrial Revolution
(1760–1840), however, smoky air left
deposits of soot on trees and
buildings in British cities, and the
black form became much commoner.
By 1895, 95 percent of peppered
moths in Britain’s cities were black,
as paler moths were eaten by birds
because their coloring provided no
camouflage. This phenomenon
continues to act as an example of Escherichia (E.) coli bacteria
Two peppered moths exhibit can cause serious gut and other
Darwin’s theory in action today, as evolution at work, the lower one an infections that will be increasingly
the pale moth becomes common example of industrial melanism. The difficult to treat as drug-resistant
once more due to the declining soot dark variety began to appear in British strains of E. coli multiply.
concentrations in Britain’s cities. ■ cities in the early 1800s.


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
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
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.
structure of DNA.
2000s Researchers in the
field of epigenetics describe
inheritance by mechanisms
other than through the DNA F2 GENERATION
sequence of genes.
1 green 3 yellows


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

dominant or recessive. When both

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

Pioneering geneticist Gregor Johann Mendel

Mendel published his paper in
1866, but no one took much notice Born Johann Mendel in 1822
until 1900, when the botanists on a farm in Silesia—then part
Hugo de Vries, Carl Erich Correns, of the Austrian Empire and
(phenotypes) that had only two and Erich Tschermak von now in the Czech Republic—
distinct forms—for example, white Seysenegg discovered his work. Mendel studied philosophy
or purple flowers. When examining Scientists then began proving and physics at the University
the trait of yellow or green peas, Mendel’s theories more widely. of Olomouc (1840–43). At this
Mendel took green pea plants and Within just ten years, scientists time, he became interested
cross-pollinated them with yellow named the pairs of factors “genes” in the work of Johann Karl
pea plants. The peas produced from and showed that they are linked on Nestler, who was researching
hereditary traits in plants
this parent generation were all yellow chromosomes. It is now known that
and animals. In 1847 Mendel
and Mendel named them the F1 inheritance is far more complex
entered a monastery, where
generation. He then cross-pollinated than Mendel recognized, but his he was given the name
pea plants from the F1 generation meticulous research continues to Gregor. He then went on
with each other to produce the F2 form the basis for modern studies. ■ to study science further at
generation. He found that some Vienna University (1851–53).
peas produced were yellow and When Mendel returned
some were green. The F1 generation to his monastery in 1853,
showed only one trait (yellow), which the abbot Cyril Napp gave
Mendel called “dominant.” However, him permission to use the
in the F2 generation 75 percent had gardens for his research into
the dominant yellow trait and 25 hybridization. Mendel himself
percent displayed the nondominant became an abbot in 1868 and
—or “recessive”—green trait. no longer had time for his
experiments. Although he
never received credit for his
Laws of inheritance discoveries during his lifetime,
Mendel theorized that every pea he is widely regarded as the
plant has two factors controlling founder of modern genetics.
each trait. When plants are cross-
pollinated, one factor is inherited Key works
from each plant. A factor can be
1866 “Experiments with Plant
Hybrids,” Verhandlungen des
Pea plants provided the raw data naturforschenden Vereines
that Mendel used to develop his in Brünn
theories explaining the transmission of
traits from one generation to the next.


Francis Crick (1916–2004),
Rosalind Franklin (1920–
58), James Watson (1928–),

Maurice Wilkins (1916–2004)
1910–29 US biochemist
Phoebus Levene describes the

chemical components of DNA.
1944 US researchers Oswald
Avery, Colin Macleod, and
Maclyn McCarty show that

DNA determines inheritance.
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.

he discovery of the structure
of DNA (deoxyribonucleic
acid) in 1953 is one of
the most important scientific
breakthroughs to date. It offered
the key to understanding the very
building blocks of life and explained
how genetic information is stored
and transferred. Englishman Francis
Crick and American James Watson
famously celebrated their joint
discovery in a low-key fashion
at their local pub in Cambridge,
followed by a letter published in
the journal Nature. Their discovery
had enormous potential for scientific
advances and had an important
impact on many fields of research,
from medicine to forensic science,
taxonomy, and agriculture. The
ramifications of their work still
reverberate today, as methods of


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

College, London, Franklin and

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

Genetic engineering useful—have greatly simplified

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


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

Good and bad errors

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


DNA barcoding
The idea of DNA barcoding
was first raised in 2003 when
a team at the University of
Guelph, Canada, suggested
that it would be possible to
identify species by analyzing
a common section of their
DNA. Led by Dr. Paul Hebert,
researchers chose a region in
the gene known as cytochrome
c oxidase 1 (“CO1”), made up
of 648 base pairs. This region
is quick to analyze, but the
sequence is still long enough
to differentiate between and
within animal species.
Different gene segments can
be used for other forms of life.
The first part of the
barcoding system involves
of disorders caused by gene Mutated blood cells occur in cataloguing samples of known
mutations include cystic fibrosis sickle-cell disease—a genetic disorder species. The DNA is extracted
and sickle-cell disease. passed on when both parents carry
and organized into a sequence
the faulty gene. It can be painful and
Although many mutations are increases the risk of serious infections. of base pairs, a process known
harmful, occasionally a mutation as “sequencing.” The sequence
will confer an advantage on an is then stored in a computer
individual, enabling it to survive in parent. Additionally, with access database, so that when a DNA
its environment better than others to such data it is possible to sample from an unknown
of the same species. This type of screen embryos for known genetic species is sequenced and
mutation may end up being passed disorders before implantation in the entered into the database, the
on through the process of natural womb. By March 2018, the DNA of computer will match it with
existing records. The barcoding
selection. Over many generations, around 15,000 organisms had been
technique has proved useful
mutation is a mechanism for sequenced. Such information can for taxonomy, helping classify
diversification, survival of the help show how animals are related animals and plants.
fittest, and ultimately evolution. in the evolutionary line and how
they have diversified.
The human genome While the discovery of the
On April 14, 2003, scientists composition and structure of DNA
completed the lengthy task of has revolutionized the science of
mapping (sequencing) the entire heredity, it is worth noting that
human genome. Geneticists worked the regions of DNA used for coding
out the precise position of all the proteins account for just 2 percent With genetic
base pairs in a chain of some three of the entire human genome. engineering, we will
billion of the base nucleotides The nature of the other 98 percent be able … to improve
comprising an estimated 30,000 is not yet fully understood by the human race.
individual genes. This has allowed geneticists, but it is believed that at Stephen Hawking
geneticists to identify new genes least some of these regions involve
and the role they play in organisms. the regulation of the way genes are
Armed with this knowledge, expressed, or activated. It seems
an individual can find out if they that many more discoveries await
have inherited a faulty gene from a future geneticists. ■



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.
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.
1982 Richard Dawkins argues Animals that warn
in The Extended Phenotype Male black widow
others of approaching
spiders mate
that the study of an organism predators sacrifice
even though the
should include analysis of themselves at the
females eat them
how its genes affect the expense of the
immediately after.
surrounding environment. wider group.

2002 Stephen Jay Gould

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


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

circumstances in which the gene

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




Robert MacArthur’s Dan Janzen observes the

Joseph Grinnell publishes research on North American interdependence of acacia
his research on the California warblers shows how different trees and the ants that reside
Thrasher, establishing the species can avoid directly on them, and concludes that
basis for the theory of competing with each other the species evolved in a
ecological niches. in order to coexist. mutualistic manner.

1917 1957 1965

1925–26 1961 1969

The Lotka-Volterra model uses Joseph Connell reveals that Robert Paine
a mathematical equation to different types of barnacle coins the term “keystone
describe the interactions between thrive in different tidal species” to describe species
predator and prey. zones, although they could, that play a crucial role in
in theory, live in any of them. ecosystem functions.

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


Research published by
Ronald Pulliam, Eric Charnov, Robert Sterner and James Elser
and Graham Pyke expands on pioneer the study of ecological
Roy Anderson and Robert the optimal foraging theory that stoichiometry—how ratios of
May demonstrate how animals try to gather resources different chemicals within
epidemics affect animal while wasting as little living organisms change
population growth rates. energy as possible. with certain reactions.

1970s 1977 2002

1972 1991

Knut Schmidt-Nielsen Earl Werner publishes

publishes How Animals his findings about
Work. The book hugely nonconsumptive
influences the field effects of predators
of ecophysiology. on prey.

laboratory projects. As William E. to play, too—as Earl Werner New technology

Odum put it in 1959, “the ecological demonstrated 30 years later. His Technological advances—including
niche of an organism depends not work revealed the non-consumptive sophisticated chemical sampling
only on where it lives, but also on impact of predatory dragonfly larvae techniques, satellites with remote
what it does.” on the behavior and physical sensing equipment, and computers
development of their tadpole prey. capable of rapidly processing huge
From field to lab Since the mid-20th century, quantities of data—have opened
Laboratory experiments and field many new ideas on ecological up new areas of study.
observations are the main methods processes have emerged. Work by Ecological stoichiometry, for
of providing data for the study Robert MacArthur and others on example, studies the flow of energy
of ecological processes, but field competition between species led to and chemical elements throughout
experiments—in which a local the development of optimal foraging food webs and ecosystems, from the
environment is manipulated to test theory, which seeks to explain why molecular level up. Like so many
a hypothesis—were not conducted animals choose to eat some food ideas in ecology, its origins can be
with scientific rigor until Joe items and not others. Mutualistic traced back many years, but only
Connell’s work with barnacles relationships became better took hold with Robert Sterner and
in Scotland. His experiments—the understood through the research James Elser’s 2003 book Ecological
results of which were published of biologists such as Daniel Janzen. stoichiometry: The biology of
in 1961—were meticulously planned Robert Paine’s work with starfish elements from molecules to the
and observed, and were repeatable. and mussels also highlighted the biosphere. New techniques such
Connell set the “gold standard” concept of keystone species— as this will undoubtedly continue
for fieldwork, but experiments in those that have a disproportionate to deepen our understanding of
laboratories still have a vital role influence on their ecosystems. processes in ecology. ■



he predator–prey equations
IN CONTEXT Populations of two species, are an early example of the
one predator, the application of mathematics
other prey, interact. to biology. Formulated in the 1920s
Alfred J. Lotka (1880–1949),
by American mathematician Alfred
Vito Volterra (1860–1940) J. Lotka and Italian mathematician
BEFORE and physicist Vito Volterra, the
1798 British economist two equations—also known as
Thomas Malthus shows that The prey has access to the Lotka–Volterra equations—
the rate at which the population food and its population describe the way in which the
growth is exponential. population of a predator species
changes increases as the size
of the population grows. and that of its prey fluctuate in
relation to each other.
1871 In Lewis Carroll’s novel Lotka proposed the equations
Through the Looking Glass, in 1910, as a way of understanding
the Red Queen tells Alice, When prey animals meet the rates of autocatalytic chemical
“you have to run just to stay a predator, they reactions—chemical processes
in the same place.” are eaten. that regulate themselves. In the
following decade, he applied
AFTER the equations to the population
1973 American biologist Leigh dynamics of wild animals.
Van Valen proposes the Red In 1926, Vito Volterra arrived
Queen effect, which describes Eating prey results at the same conclusions. He had
the constant “arms race” in more predators. become interested in the subject
between predators and prey. after meeting Italian marine
biologist Umberto D’Ancona.
1989 The Arditi–Ginzburg D’Ancona told Volterra how the
equations offer another model percentage of predatory fish
of predator–prey dynamics caught in nets in the Adriatic
More predators
by including the impact of the Sea had greatly increased during
results in less
ratio between predator and prey. World War I. This change was
prey, reducing the
number of predators. clearly linked to the drastic
reduction in fishing during the

Vito Volterra Born in 1860 in Ancona, Italy, the Volterra refused to swear loyalty
son of a Jewish cloth merchant, to Italy’s fascist dictator Benito
Vito Volterra grew up in poverty. Mussolini and was dismissed
Despite this, in 1883, aged just 23, from the University of Rome.
he secured a position as professor Forced to work abroad, he only
of mechanics at the University returned to Italy for a short time
of Pisa and began a career as before his death in 1940.
a mathematician. Further
professorships at the universities Key works
of Turin and Rome followed. In
1900, Volterra married, fathering 1926 “Fluctuations in the
six children, although only four Abundance of a Species
survived to adulthood. He was Considered Mathematically,”
made a senator of the Kingdom Nature
of Italy in 1905 and worked on the 1935 Les associations
development of military airships biologiques au point de vue
during World War I. In 1931, mathématique

See also: Evolution by natural selection 24–31 ■ The selfish gene 38–39 ■ Ecological niches 50–51 ■ Competitive exclusion
principle 52–53 ■ Mutualisms 56–59 ■ Keystone species 60–65 ■ Optimal foraging theory 66–67

A cheetah pursues a Thomson’s increases as the population grows. occurring factors. As a result, wild
gazelle. The predator–prey equations From this theory, Malthus predicted populations should in theory be
are able to model the way populations a catastrophic future for humanity. more or less static, fluctuating only
of both species will change in response
to the activities of the other.
The number of humans was around the carrying capacity,
growing much more quickly than assuming the random impacts of
the amount of food that could be catastrophic events are ignored.
war years, but D’Ancona could not produced by the world’s farmlands. However, this relative
explain why less fishing did not Eventually, Malthus argued, a point equilibrium did not always match
produce more fish of all kinds in the would be reached when the human up with observations—as in ❯❯
nets. Using the same equations as population would succumb to
Lotka, Volterra eventually explained global famine and decline.
the fluctuations in both the predator Malthus’s bleak vision did not
and the prey species. happen, thanks to technological
advances in agriculture and the
Population principles development of artificial fertilizers,
At the time Lotka and Volterra but his population model became The food species
made their calculations, the science applicable to species populations cannot, therefore, be
of population dynamics was still within ecosystems. Every habitat, exterminated by the
in its infancy, having barely moved and the niche occupied by a species predatory species, under
on since the population studies of within its community of organisms, the conditions to which
British economist Thomas Malthus has a carrying capacity—the our equations refer.
in the late 18th century. According maximum population that can Alfred J. Lotka
to Malthus’s theory, a population be supported by the resources
grows or declines rapidly as long available, such as water, space,
as the environmental factors for food, and light. Any rise in
survival are constant, and the rate population above this level is
at which that population changes likely to be reduced by naturally

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.

Volterra was able to show that the

average sizes of predator and prey
populations do indeed oscillate but
the rate at which each population
is growing or declining is always
changing and almost never
matches the changes experienced
by the other population. To
eliminate variables, Volterra made
a series of assumptions: first, that
the prey and predator species have TIME

as antelopes and deer—and
mammalian carnivores, like the
big cats and wolves, is an example
of this evolutionary arms race. The
hoofed animals have long legs,
extended by walking on the very
tips of thickened and fused toe
bones. This adaptation allows
them to outrun and outjump their
predators. In response, big cats—
such as lions and tigers—have
evolved speed and strength to
bring down large, fleet-footed prey
in surprise attacks. Wolves have
evolved the stamina to run for
long distances without stopping.
This allows them to work as a
team to chase down their prey
and kill them when the exhausted
prey collapse.
While the predator–prey
equations offer an insight into
the population dynamics of two
species, the assumptions they rely
on are rarely reflected in real life.
Some predators do specialize in
killing a single prey species, but
other factors in the ecosystem
also affect their populations.

Other applications
The Lotka–Volterra equations have
been used to study the dynamics of another species but also the prey The parasitoid wasp lays its eggs
food chains and food webs in which species of a third. They have in aphids (the smaller, yellow insects
one species may be a predator of also been used to examine the shown above). It is called a parasitoid
because the wasp’s larvae later eat the
relationship between host and aphids as they grow.
parasite species, which bears
some resemblance to that between
prey and predator. Parasites often thanks to beneficial genes, certain
specialize in one host species— individuals in a host population
Volterra was interested a relationship that should resemble are able to maintain their fitness
in a mathematical the one described by the Lotka– despite the attacks from parasites.
theory of ‘the survival Volterra equations. However, in The parasites constantly evolve to
practice the process of evolution exploit these seemingly immune
of the fittest.’
is thought to interfere with this. individuals, and therefore the
Alexander Weinstein A parasite does not usually kill beneficial genes in the host
Russian mathematician
its host (those that do are called population also change. In this way,
parasitoids), but can reduce its evolution is happening all the time,
fitness. The Red Queen evolutionary as the parasite and host battle it
theory, proposed in the 1970s by out—although everything appears
Leigh Van Valen, describes how, to stay the same. ■



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
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
1968 A study by Australian Existence of each
D.R. Klein of the introduction, species is Finding a unique niche
increase, and die-off of reindeer determined by a is the circumstance that
on St. Matthew Island, Alaska, slender thread removes competition.
identifies the destructive niche. of circumstances.

See also: Competitive exclusion principle 52–53 ■ Field experiments 54–55 ■ Optimal foraging theory 66–67
■ Animal ecology 106–113 ■ Niche construction 188–189

An ultra-specialist Giant pandas occupy a very Pandas eat different parts of

specialized ecological niche, as bamboo plants according to the
their diet consists mainly of seasons. In late spring, they
bamboo. Bamboo is a poor food prefer the first green shoots.
source, low in protein and high in They eat leaves at other times
cellulose. Pandas can digest only of the year, and stems in winter
a small proportion of what they when little else is available.
eat, which means they have to Pandas have evolved muscular
eat a lot of bamboo—as much as jaws and a pseudothumb to
28 lb (12.5 kg) each day—and manipulate bamboo stems. Their
forage for up to 14 hours a day. digestive tract is inefficient at
It is unclear why pandas have processing large quantities of
become so dependent on bamboo, plant material because it
but some zoologists suggest it is remains similar to that of its
because it is an abundant and carnivorous ancestors, although
reliable food source, and pandas digestion is helped by the
are not skilled predators. bacterial fauna in their gut.

habitat known as chaparral, and factors. Thirty years later, George coexist (niche partitioning), and
how it escaped predators by Evelyn Hutchinson expanded the the overlap of resources by different
running through the underbrush. definition yet further. He argued animals and plants (niche overlap).
The thrasher’s camouflage, short that a niche should take into
wings, and strong legs were account all of an organism’s The importance of habitat
perfectly adapted for life in this interactions with other organisms Ecological niches depend on the
environment. Grinnell saw the and its nonliving environment, existence of a stable habitat; small
chaparral habitat as the thrasher’s including geology, acidity of soil or changes can eradicate niches that
“niche.” His idea also allowed for water, nutrient flows, and climate. organisms once filled. For example,
“ecological equivalence” in plants Hutchinson’s work encouraged dragonfly larvae only develop
and animals, whereby species others to explain the variety of within a certain range of water
distantly related and living far resources used by a single acidity, chemical composition,
apart could show similar organism (niche breadth), the ways temperature, and prey, and with
adaptations, such as feeding in which competing species a limited number of predators.
habits, in similar niches. In the The right vegetation is needed by
Australian outback, for instance, adult females for egg-laying, and
babbler bird species forage in the by larvae for metamorphosis.
scrubby vegetation in a similar way The dragonfly also impacts its
to the unrelated thrasher. Grinnell environment: its eggs are food for
also identified “vacant” niches— amphibians; its larvae, which are
habitats that a species could [A niche] is a highly both predators and prey, add
potentially occupy, but where it abstract multi- nutrients to the water; and the
was not present. dimensional hyperspace. adults prey on insects. These
George Evelyn requirements and impacts define
Widening the niche Hutchinson its ecological niche. Hutchinson
In the 1920s, ecologist Charles argued that for a species to persist,
Elton looked beyond a simple conditions had to be within the
habitat definition for “niche.” For required ranges. If conditions moved
him, what an animal ate and what outside the niche requirements, a
it was eaten by were the primary species could face extinction. ■



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
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
1959 G. Evelyn Hutchinson
extends Gause’s ideas and Cape May Blackburnian Black-throated
produces a ratio describing the Warbler Warbler Green Warbler
limit of similarity between two
competing species.
1967 Robert MacArthur and
Richard Levins use probability Five species of
theory and Lotka–Volterra warblers are able to
equations to describe how share the same tree,
coexisting species interact. because each inhabits
its own “niche.” Living
in this way, without
Bay-breasted Yellow-rumped much overlap, the
Warbler Warbler birds do not compete.

See also: Evolution by natural selection 24–31 ■ Ecological niches 50–51 ■ Animal ecology 106–113 ■ The ecosystem
134–137 ■ The ecological guild 176–177 ■ Niche construction 188–189 ■ Invasive species 270–273

genetics. In fact, the competitive

exclusion principle—although a
useful theoretical model—is rarely
seen in nature, simply because, in a
bid to survive, a weaker competitor
tends to quickly move on or adapt. Let us make for this purpose
an artificial microcosm… let
Avoiding competition us fill a test-tube with nutritive
Most creatures can make the medium and introduce to it
changes necessary for survival. A several species of protozoa
variety of birds can live in a garden consuming the same food
during any one year because they or devouring each other.
all operate in different “niches.” Georgy Gause
They have contrasting beak shapes
and sizes that allow them to eat
The red squirrel is smaller than the
gray, and has a more restricted diet different types of food—the robin
and habitat. Reds may also die from the preferring insects, the finch eating
squirrel parapoxvirus, which is carried seeds. Their choice of habitat and
by the grays but does not affect them. feeding times might also vary; this
is known as resource partitioning. at different heights and depths of
In 1957, Robert MacArthur the foliage. In this way they avoid
nature, he proposed, there were too noted this phenomenon in North competing with each other.
many variables to draw conclusions American warblers. The five
about how ecological mechanisms species he observed, each with An invasive competitor
work. He argued that little progress distinctive, colorful markings, Problems often arise if an exotic
had been made since Darwin’s era flitted in and out of coniferous species is suddenly introduced
in understanding how species trees, feeding on bugs and other to an ecosystem. Britain’s red
compete for survival, whereas the insects. They could coexist in one and gray squirrels provide a clear
experimental method had produced habitat because they did not try to example. When the grey arrived
great advances in areas such as feed in the same part of the tree but from America in the 1870s, both
squirrel species competed for the
Types of competition same food and habitat, which put
the native red squirrel populations
The Competitive Exclusion “limiting resource,” the one that under pressure. The gray had
Principle covers two main types both require in order to breed. the edge because it can adapt
of competition. Intraspecific Ecologists make a further two its diet; it is able, for instance, to
competition is between distinctions. Interference is eat green acorns, while the red can
individuals of the same species when two organisms fight only digest mature acorns. Within
and ensures the survival of directly with each other over a the same area of forest, gray
the fittest, so that only the limited resource, such as a mate squirrels can decimate the food
healthiest individuals—or those or a preferred food. Exploitation supply before red squirrels even
best adapted to a particular is indirect competiton, such as have a nibble. Grays can also live
environment—will breed. The stripping out a resource so there more densely and in varied
second type is interspecific: is none left for the competitor;
habitats, so have survived more
competition between two this can be seen in plants, when
different species that rely on a species’ uptake of nutrients or easily when woodland has been
the same resources. The most water is more efficient than that destroyed. As a result, the red
important of these will be the of its neighbors. squirrel has come close to
extinction in England. ■



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.
behave the way they do would be A laboratory, however, is an
Joseph Connell (1923–)
largely speculative. Rigorous artificial environment, where
BEFORE observation is also essential, but, organisms may not behave as they
1856 British scientists John much of the time, experimentation do in their natural habitat. For
Lawes and Joseph Gilbert start is needed for a full understanding example, bats leaving a roost at
the Park Grass Experiment at of those observations. dusk may follow different routes
Rothamsted, to test how Three main types of ecological to their foraging areas in spring
different fertilizers affect the experiments are used to test and late summer. The potential
yield of hay meadows. theories: mathematical models, reasons for the switch—changes
laboratory experiments, and field in prey distribution and predator
1938 Harry Hatton, a French experiments. Each method has its threats; seasonal differences in
ecologist, conducts one of merits, but it is only recently that tree cover; or human disturbance
the first marine ecology field the benefits of field experiments and light pollution—cannot
experiments, on barnacles be established in a laboratory.
on the Brittany coast. Mathematical modeling might help
predict patterns, but would be less
AFTER effective at identifying the causes
1966 American ecologist of change. To understand the bats’
Robert Paine removes the behavior, a study of their natural
starfish Pisaster ochraceus from environment is crucial, and
tide pools in a Pacific coast this is achieved only through
ecosystem, to test the effect research in the field.
of its absence on other species. Field experiments allow different
factors to be manipulated to test
1968 The Experimental Lakes their relevance. In the bat example,
Area, comprising 58 freshwater
lakes, is established in Ontario,
Canada, to study the effects Rain forest ecosystems are some of
the most species-rich environments
of nutrient enrichment
on Earth. This makes them especially
(eutrophication). valuable sites for ecologists to conduct
experiments in the field.

See also: Ecological niches 50–51 ■ Modern view of diversity 90–91 ■ Animal behavior 116–117 ■ The ecosystem 134–137
■ Niche construction 188–189

street lights could be switched off to Joseph Connell’s barnacle experiment

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



Dan Janzen (1939–)


1862 Charles Darwin proposes
that an African orchid with


a long nectar receptacle must
be pollinated by a moth with


an equally long proboscis.
1873 Belgian zoologist Pierre-
Joseph van Beneden first
MUTUALISMS uses the term “mutualism”
in a biological context.
1964 The term “coevolution”
is first used by American
biologists Paul Ehrlich and
Peter Raven to describe
the mutualistic relations
between butterflies and
their food plants.
2014 Researchers discover
an unusual yet beneficial
three-way mutualism involving
sloths, algae, and moths.

n biology, there are several
kinds of interaction between
organisms. One species in an
ecosystem may lose out to another
when competing for the same
resources. A prey species may be
eaten by a predator. There are also
symbiotic relationships, in which
one species benefits but not at the
expense of the other, or where one
organism does not benefit but still
survives. In the relationship known
as “mutualism,” both organisms
benefit from the relationship.

A tree and its ants

In the mid-1960s, Daniel Janzen,
a young American ecologist,
became fascinated by the amazing

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


1. Adult female wasp

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

and thence disperse their spores.

3. Inside mature fig,
When a bird swallows a fruit, it
male wasps fertilize carries the seeds with it
new females and dig as it flies away; the indigestible
escape tunnels for them seeds may be excreted in faeces
far from where they were eaten.
The fig wasp and the fig share a complex In all these situations, the plants
service-resource mutualism, in which the wasp provides provide a resource (food) and the
the service of pollination and the fig plants provide mammals, flies, and birds provide
a secure environment for the wasp eggs to develop. a service (transport).
However, not all mutualistic
acacia would have no means of the animal. It is estimated that relationships involve plants. In
defending itself. And if the acacia nearly three-quarters of flowering Africa, birds named oxpeckers
trees were removed, the ants would plants (some 170,000 species) are and herbivorous mammals such
have no home. pollinated by 200,000 animal as impalas and zebras practise
species. Typically, a pollinating another kind of service-resource
Benefits for all insect is attracted to a flower by its mutualism. The oxpeckers pick
There are two fundamental types colours or scent to drink nectar or ticks from the mammals’ fur,
of mutualism—service-resource collect pollen, and pollen attaches removing irritation and a source
and service-service relationships. to part of the insect’s body to be of disease, while at the same time
They are defined by the nature carried to the next flower, where having a meal. Oxpeckers also
of the relationship between the it is deposited. The flower and its make loud calls when they sense
partner organisms, whether it is pollinator have evolved to make danger, alerting the mammal host
the provision of a service or the this mechanism work effectively. as well as other oxpeckers.
supply of a resource—both are Some plants have also evolved In the insect world, some ants
usually key to survival. Service- a service-resource relationship in and aphids carry out a different
resource relationships are common which birds and mammals disperse form of service-resource mutualism.
in nature, with the fertilization, or their seeds, spores, or fruit. Seeds While the aphids feed on plants,
pollination, of flowers by butterflies, may become attached to the fur of the ants protect the aphids.
moths, bees, flies, wasps, beetles, a mammal browsing the plant’s Subsequently, the ants consume
bats, or birds the most widespread leaves; when the mammal wanders the honeydew that the aphids
example. The resource (pollen) is away, it disperses the seed. The release, using a “milking” process
provided by the flower, and the vile odor of stinkhorn fungi attracts on their smaller partners, by
service (pollination) is provided by flies, which lick the fungi’s slime stroking them with their antennae.

Service-service mutualisms, in return for the protection offered of orchids. Like many other
which both organisms offer each by the sea anemones’ venomous flowering plants, orchids rely on
other protection, are far less tentacles, the clownfish deters insects to pollinate them. Some
common. One unusual relationship predatory butterfly fish, removes have extraordinary structures in
takes place in the western Pacific parasites from its host, and also which to hold nectar and pollen.
Ocean, between around 30 species provides nutrients from its faeces. To lure the insect pollinators, the
of clownfish and 10 species of plants offer them a drink of energy-
venomous sea anemones. The sea Cooperative evolution giving nectar. This fascinated
anemones’ stinging, toxin-filled Relationships between service and Darwin, who was given a specimen
nematocysts, or capsules, on their resource providers have developed of the Madagascar orchid in 1862.
tentacles kill most small fish that over millions of years in a process The flower stores its nectar in a
come close, but not the clownfish. called “coevolution”—the evolution hollow spur nearly 30 cm (12 in)
Its thick layer of protective mucus of two or more species that affect long. Darwin and Wallace
provides immunity against the each other reciprocally. speculated that only a large moth
anemone’s sting, allowing the fish The term coevolution was could have a proboscis long enough
to live within the tentacles. In coined by American biologists Paul to reach the nectar—a theory
Ehrlich and Peter Raven in 1964, eventually proven in 1997. If the
but a century before the word orchid’s spur were shorter, a moth
The clownfish and sea anemone
could both survive without the other’s existed, the naturalists Charles could drink without picking up
protection, but their coevolved mutual Darwin and Alfred Russel Wallace pollen and so would not pollinate
relationship gives them a much higher were already aware of the concept, the flower. If the spur were longer,
chance of survival. not least through their observation a moth would not visit. ■



keystone species plays
IN CONTEXT a crucial role in the way
an ecosystem functions,
even though it is often a small
Robert Paine (1933–2016)
part of the overall biomass of the
BEFORE ecosystem. Because it exerts a Do you want an auto
1950s In Kenya, farmer and disproportionately large effect on mechanic who…can name,
conservationist David the environment relative to its list, and count all of the parts
Sheldrick introduces elephants biomass, if a keystone species of your engine, or one who
to Tsavo East National Park, disappears from an ecosystem, really understands how each
and discovers this results in a that ecosystem will change part interacts with the others
major increase in biodiversity. dramatically. The importance of to make a working engine?
keystone species was brought to Robert Paine
1961 Fieldwork by American light by the American biologist
ecologist Joseph Connell on Robert Paine—who derived the
Scotland’s rocky shores shows term from the central “keystone” at
that removing predatory the top of an arch that stops it from
whelks alters the distribution collapsing—in his 1969 article “A
of their barnacle prey. Note on Trophic Complexity and
Community Stability.” clear impact on many others. Paine
AFTER developed the idea to include the
1994 In the US, a group of The keystone concept concept of “trophic cascades”—the
ecologists led by Brian Miller In the 1960s, Paine spent several strong, top-down effects that ripple
publishes a paper explaining years studying the animals of the through an ecosystem and its
the valuable role prairie dogs intertidal zone of Tatoosh Island organisms. Since Paine’s work
play as a keystone species. on the Pacific coast of Washington with starfish, several studies
State. He removed the ocher have demonstrated that there are
2016 Fieldwork leads marine starfish and watched its key prey,
ecologist Sarah Gravem to a mussel whose numbers had
conclude that organisms can Black-tailed prairie dogs look
been kept in check by the starfish, out from their burrow in a field in
be keystone species in some dominate the zone, replacing other Wyoming. Study of this species has
places but not in others. subordinate species. The removal revealed its key role in fostering
of a single, keystone species had a diversity in its native habitat.

See also: Predator–prey equations 44–49 ■ Mutualisms 56–59 ■ Animal ecology
106–113 ■ Trophic cascades 140–143 ■ Evolutionarily stable state 154–155

Whelks feed on They also display

barnacles; they aggression toward
are predators. their prey.

Robert Paine
In areas where
Whelks are like there are large Born in 1933, in Cambridge,
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 ❯❯

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.

Keystone predators
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.
Since 1911, legal protection has led
to a slow increase in numbers.
Sea otters are important
because they eat large numbers
of sea urchins. These seafloor-
dwelling invertebrates graze on the
lower stems of kelp that grow up

Every species in the coastal

zone is influenced in one way
Bechler Snake River
or another by the ecological
effects of sea otters.
James Estes
American marine biologist
Each pack of wolves in the
Yellowstone National Park has its own
SCALE territory. Many of the territories overlap,
and numbers fluctuate from year to
10km (6 miles) year, with 108 wolves recorded in 2016.


Reintroducing Beavers were wiped out in the UK several dams on the headwaters
beavers to the UK 400 years ago, but the beneficial of the Tamar River, creating 13
role of this keystone mammal is new freshwater pools and making
now better understood. Beavers surrounding areas wetter.
are ecological engineers, building In Devon, the damp areas
dams and canals, and their created by beavers led to an
presence increases biodiversity. increase in the number of
In 2009, 11 beavers were bryophyte species (mosses and
reintroduced to Knapdale Forest, liverworts), and the range of
Scotland, and in 2011, the Devon aquatic invertebrates has risen
Wildlife Trust introduced a pair to from 14 to 41 species. Increased
a fenced enclosure. Both projects numbers of flying insects have
have been monitored to test their also improved bat diversity, with
impact on the environment. In two nationally rare bat species
Knapdale Forest, the beavers’ drawn into the area. More
dams changed the water level of beaver reintroduction projects
a loch, and Devon’s beavers built are now planned in the UK.

from the seabed, causing it to drift herbivores, such as beavers. Within fruited plant species share one or
away and die. If the kelp disappears, 10 years, the number of beaver two peaks of ripening each year.
however, so do the many other colonies increased from one to nine. Fig trees bear fruit throughout the
marine invertebrates that graze on Beaver dams helped revive wetlands, year, supporting many animals
it. “Forests” of kelp also absorb large and wetland wildlife flourished. The when other trees are fruitless.
amounts of atmospheric carbon increase in elk carcasses also More than 10 percent of the world’s
dioxide and, by slowing water benefited carrion-eaters—especially bird species and 6 percent of
currents, help protect coastlines coyotes, red foxes, grizzly bears, mammals (a total of 1,274 species)
from storm surges. The protection Golden Eagles, Ravens, and Black- are known to eat figs, as do a small
that sea otters offer kelp along billed Magpies—as well as several number of reptiles and even fish.
stretches of open coast is therefore smaller scavengers. Fig trees therefore provide a vital
particularly significant. Jaguars are apex predators in support mechanism for fruit-eating
Unlike the sea otter, some South and Central American forests, species. Without them, fruit bats,
keystone species are also “apex” preying on more than 85 species. birds, and other creatures would
predators at the top of the food Although there are very few jaguars decline or disappear. ■
chain, such as the gray wolf. Before in any given area, their impact on
1995, there had been no gray wolves the numbers of other predators—
in Yellowstone National Park for at such as caimans, snakes, large
least 70 years. American elk were fish, and large birds—as well as
common in the park, but there was herbivores, such as capybaras and
just a single colony of beavers. That deer, has a significant ripple-down
year, 31 wolves were introduced to effect on their ecosystem. Left By protecting a keystone
the park and by 2001 their numbers unchecked, the herbivores could species such as the prairie
had increased to more than 100, devour most of the plants and dog, the public could be
largely due to the abundance of destroy the habitat on which so educated about the value of
elk for food. many other species depend. ecosystem conservation.
The presence of wolves in the Brian Miller
park forced the elk to become more Keystone plants American ecologist
mobile. Rather than over-grazing Not all keystone species are
willow, aspen, and cottonwood trees animals. One example is the fig
in favored locations, the elk moved tree, of which there are about 750
on, allowing plants to regenerate species, mostly found in tropical
and provide a food source for other forests. In this habitat, most fleshy-



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.
their nutrients and water from soil,
Ronald Pulliam (1945–),
and sunlight provides the energy Foraging theories
Graham Pyke (1948–), and
for photosynthesis. Animals The first theory of foraging by
Eric Charnov (1947–) generally have to work harder to wild animals did not emerge until
BEFORE find their food—they have to move, the mid-1960s, when Americans
1966 John Merritt Emlen, and this uses extra resources. Robert MacArthur and Eric Pianka
Robert MacArthur, and Eric Optimal foraging theory (OFT) examined the question of why,
Pianka outline the concept proposes that animals try to gather when a range of food was available
of optimal foraging in two resources in the most efficient way to them, animals often restricted
articles published in the to avoid using additional energy. themselves to a few preferred types
Searching for and capturing food of prey. They argued that natural
American Naturalist magazine.
takes energy and time. The animal selection favored animals whose
AFTER needs to gain maximum benefit behavior maximized their net
1984 Argentinian–British for minimal effort in order to energy intake per unit of time spent
zoologist Alejandro Kacelnik achieve optimal fitness. OFT foraging. An animal’s foraging time
researches the foraging includes searching for prey and the
behavior of starlings to killing and eating of the food
illustrate the marginal (handling time).
value theorem (MVT). These ideas were developed by
American ecologists Ronald Pulliam
1986 Belgian ecologist Patrick and Eric Charnov and Australian
Meire investigates prey Diets should be broad ecologist Graham Pyke. It seems
selection by oystercatchers. when prey are scarce, that OFT works best for mobile
but narrow if food foragers seeking immobile prey, and
1989 Swiss environmental is abundant. some researchers believe it is less
scientists T. J. Wolfe and Paul Eric Pianka relevant when prey are mobile.
Schmid-Hempel examine how
the weight of nectar carried Key choices
by bees has an effect on the Animals must choose which types
bees’ foraging behavior. of food to eat, which is rarely
straightforward. For example,

See also: Evolution by natural selection 24–31 ■ Predator–prey equations
44–49 ■ Competitive exclusion principle 52–53 ■ Mutualisms 56–59

small clams was better spent

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



Roy Anderson (1947–),

Robert May (1936–)
1662 English statistician John

Graunt seeks to classify causes
of death in London in Natural

and Political Observations
made upon the Bills of Mortality.
1927 Scottish scientists

Anderson Gray McKendrick
and William Ogilvy Kermack
develop an epidemic model for
infected, uninfected, and
immune individuals.
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.

pidemiology is the study
of how disease spreads
through a population. Its
initial application was to human
diseases, but its methods have
been recognized as an effective
way of modeling populations of
other organisms, too.
Ecologists have long known
that the size of an animal or plant
population and its growth rate
depend on the availability of food,
living space, and levels of
predation. In the 1970s, British
epidemiologist Roy Anderson and
Australian scientist Robert May
showed how parasites and
infections from pathogens such as
bacteria and viruses limited the

See also: The microbiological environment 84–85 ■ Microbiology 102–103 ■ The ubiquity of mycorrhizae 104–105
■ Biodiversity and ecosystem functioning 156–157

Map of deaths from cholera in London in 1854

1–4 deaths Street

5–9 deaths Soho


10–15 deaths

Broad Street

d Str
B r oa
t re

Fatalities in London’s et

cholera outbreak of Golden re

1854 were linked to the Square er
central pump; its water ew
was found to have
been contaminated
with infected sewage
from a stricken family.

size of a population. In wild sheep, thought to be caused by miasma—

for instance, the chief cause of a sort of poisonous vapor in the
death is lungworms, while most air—that spread from the bodies of
wild birds die from viral infections. the dead and dying. Snow was not
In ecology, the effects of disease the first to question this theory, but
have wider implications. Up to he was especially suspicious of it in
40 percent of ocean bacteria are the case of cholera.
killed each day by viruses. This In 1854, Snow plotted every
causes a “viral shunt,” because case of cholera on a map of Soho,
nutrients that would otherwise flow and found that afflicted households
up the food chain to consumers collected their water from a pump
revert to the bottom of the chain. on Broad Street (later renamed
Broadwick). He shut down the ❯❯
Human beginnings
Epidemiology has its beginnings
British doctor John Snow fought the
in the work of physician John Snow, establishment to gain acceptance for
who witnessed a cholera epidemic his belief that cholera was waterborne.
in the Soho district of London in The medical journal The Lancet finally
1854. At the time, disease was conceded that he was right in 1866.

The role of drought pump, and the epidemic soon
in plant diseases ended. This showed that cholera
was a waterborne disease that
Like other disease-causing humans contracted through
agents, a plant pathogen contaminated food and drink. A
(disease-causing agent)
decade later, Louis Pasteur’s “germ
needs a supply of susceptible
individuals to infect. Periods theory” proposed that diseases, as
of drought slow the rate of well as general rotting and decay,
plant reproduction and were the work of microorganisms.
growth, thereby reducing the
prevalence of disease. Disease model
Aridity, however, also In their 1970s studies, Anderson
weakens plants and makes and May focused first on building
them susceptible to pathogens a mathematical model to show
that thrive in dry conditions. how a microorganism could affect
These include various forms a population. This led to a set of
of fungi that attack the leaves equations that they hoped would
of grain crops, legumes, and help explain the real-life impact of A ravaged tree in North Yorkshire,
fruits. These fungi are adapted different kinds of pathogens, from UK, shows the effects of Dutch elm
to survive in a dormant state disease, a fungus spread by elm bark
bacteria and viruses to parasitic
as hardened microscopic beetles accidentally introduced to
worms and insect larvae.
bodies in soil. They can exist Europe and America from Asia.
for many years in dry soil, but In their model, a number of mice
when the soil becomes wet, were divided into three groups:
the fungi must find a host susceptible (uninfected) mice, of disease, the total would remain
within a few weeks or die. infected mice, and mice that had more or less the same, with the
They do not necessarily kill survived infection and were now rate of added mice balancing that
their host. Recent research immune. Unlike many earlier at which other mice died.
into chickpeas suggests that epidemiological models, the total For simplicity, the model
although infections from such population was not a fixed number; assumed that the diseases were
fungi do increase during a dry mice could be added either by transmitted by contact between
spell, the mortality rate of the reproduction or by additions from infected and uninfected mice. Not
affected plants goes down other populations. Mice also died all infected mice would die, so the
during a drought. from natural causes. In the absence model also included a recovery rate.
Mice that recovered would be
immune, at least initially. Immunity
to viruses is more or less lifelong,
but it is possible to become
susceptible again to the same
bacterial infection as time passes.
Sensibly used, Therefore, the calculations also
mathematical models are included a rate of loss of immunity.
no more and no less than Putting all this together,
tools for thinking about Anderson and May produced a
things in a precise way. set of equations to predict the rate
Roy Anderson and of population change in the three
Robert May initial groups of uninfected but
A summer drought produces susceptible mice, infected mice,
only sparse growth of young barley and the immune survivors. These
plants. Lack of moisture and too
much heat reduce their resistance equations could be added together
to fungi that attack their roots. to give the rate of change for the
total mouse population.

effects of hypothetical values. They by the disease. Infection numbers
found, for instance, that when the rise sharply to a maximum, then
rate of added mice was highest, the drop away. Epidemics also occur
disease had the greatest impact on when a disease is not particularly
population numbers. This suggests deadly but slows the population
Diseases such as that species with high reproductive growth rate; this has occurred with
measles and rubella, rates (introducing large numbers human diseases such as measles
with short infections of uninfected offspring) are most and chickenpox.
and lasting immunity, likely to have endemic diseases
will tend to exhibit within the population, and show Applying the theory
epidemic patterns. depressed numbers compared with The characteristics of disease
Roy Anderson species that breed more slowly. and its effects on animal and
They also explored the differing plant populations are of increasing
effects on populations of diseases ecological importance. Food
of different intensities. producers, for example, benefit from
Unlike endemic diseases, studies into the nature of parasites
in which the population’s level and the dynamics of diseases that
of infection remains consistent, can affect crops and livestock.
From their calculations, they epidemics appear in populations Conservationists also employ
deduced that a disease will persist when the growth rate of all infected epidemiology to predict how exotic
in a population whose equilibrium and uninfected members is low diseases and invasive parasites
point (the rate of new additions, compared to the death rate caused might affect fragile ecosystems. ■
balanced by the natural death rate)
is greater than the combined
effects of natural mortality, disease Venn diagram of ecological epidemiology
deaths, recovery, and transmission
Susceptible host
rate. While the disease is present,
that equilibrium point will be lower
than if the population were disease
free. If, however, the equilibrium
point of a population affected by
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.
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.



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
simple bacteria to complex scientists understand how the
Knut Schmidt-Nielsen
mammals, are adapted by natural stresses created by climate change
selection to survive in a particular impact on both wild ecosystems
BEFORE niche and habitat. Ecophysiology— and cultivated environments.
1845 The explorer Alexander for which Knut Schmidt-Nielsen’s
von Humboldt reveals that book Animal Physiology (1960) was Managing temperature
plants facing similar ecological a vital inspiration—is the study of Ecophysiology has revealed a
factors also have many an organism’s anatomy and how it number of specific adaptations
analogous features. functions (its physiology), as well as for different environments. For
how these characteristics relate to example, animals that live in colder
1859 Charles Darwin argues the challenges posed by its regions tend to have larger bodies
that organisms evolve because environment. It shows how the and smaller legs, ears, and tails
they are adapting to changed anatomy of an animal or plant is than related species living in
ecological conditions. linked to its ability to survive, and warmer climes. A larger body has
a smaller surface-area-to-mass
AFTER ratio, and therefore loses heat more
1966 Australian biochemists slowly, while smaller appendages
Marshall Hatch and Charles reduce exposure to frostbite.
Slack explain that the most In the most extreme cold, the
widespread plants are the feet of a warm-blooded animal
ones that photosynthesize From a physiological are at risk of becoming frozen to
most efficiently. viewpoint, freshwater the ground. Mammals in Arctic
is no more freely regions such as musk oxen and
1984 Peter Wheeler, a British available in the sea polar bears are adapted for life in
scientist, suggests that human than in the desert. these conditions by having thick
bipedalism—the ability to Knut Schmidt-Nielsen hairs to insulate their feet.
walk on two legs—evolved as In the Antarctic, the undersides
a thermoregulatory adaptation of penguins’ feet are insulated
that reduces the body’s by a thick layer of fat. Penguins
exposure to direct sunlight. also have a heat-exchange (or
counter-current) mechanism in

See also: Evolution by natural selection 24–31 ■ Ecological niches 50–51
■ Competitive exclusion principle 52–53 ■ Ecological stoichiometry 74–75

their legs. The warm blood arriving cooler than the air outside, so the
from the body is cooled to near moisture it carries condenses in
32°F (0°C) by the chilled blood the nose. This creates the cool,
arriving from the feet, which damp conditions needed to chill
warms to body temperature in the next in-breath.
the process.
Gazelles in Africa use a similar Future challenges
counter-current system to cool their Today ecophysiology is becoming
body temperature. They are able to increasingly focused on plants,
chill the blood entering their head, fungi, and microbes. Like animals,
giving them an advantage over they have to adapt to survive—and Knut Schmidt-Nielsen
their predators, who often overheat. studying them offers the possibility
Camels have a heat-exchange of vital discoveries for commercial Knut Schmidt-Nielsen grew
system in their nasal cavity, which and conservation purposes. ■ up in the Norwegian town of
reduces the amount of water lost in Trondheim. His interest in the
their breath. Hot, dry air is inhaled Emperor penguins survive freezing way animal physiology related
and mixes with moisture inside Antarctic temperatures thanks in part to habitat was inherited from
the nose before traveling to the to the way their bodies have evolved his grandfather who, years
lungs. The exhaled air is much to adapt to the harsh environment. before Knut’s birth, had
released thousands of flounder
(a marine fish) hatchlings into
a freshwater lake. Although
the fish thrived, they were
unable to breed because their
reproductive physiology was
adapted for life in salt water.
Schmidt-Nielsen joined
Duke University, North
Carolina, in 1954. He built a
climate-controlled space for
keeping desert animals, where
he considered the anatomy
of camels, gerbils, and other
species able to live for long
periods without water. He also
investigated the respiratory
systems of birds and the
buoyancy of fish. His 1960
textbook Animal Physiology
is still a classic work.

Key works

1960 Animal Physiology

1964 Desert Animals
1972 How Animals Work
1984 Scaling
1998 The Camel’s Nose:
Memoirs of a Curious Scientist



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
made up of chemical elements biologists Robert Sterner and James
Robert Sterner (1958–),
in varying ratios. Ecological Elser; in Ecological Stoichiometry
James Elser (1959–)
stoichiometry considers the (2002), they used mathematical
BEFORE balance of these elements, and models to demonstrate the
1840 German biologist and how the ratios change during application at every level, from
chemist Justus von Liebig chemical reactions. Studying molecules and cells to individual
asserts that the limitations such ratios throws light on the plants and animals, populations,
on agriculture productivity are way the living world operates, communities, and ecosystems.
primarily chemical. revealing how organisms obtain
the nutrients and other chemicals Key chemicals
1934 US oceanographer Alfred they require for life from the In ecological research, the three
Redfield measures the atomic resources in their environment. main elements examined are
ratio of carbon, nitrogen, and carbon (C), nitrogen (N), and
phosphorus (C:N:P) in plankton phosphorus (P), because each
and seawater, and finds it to be plays a vital role. Carbon is a basic
relatively consistent in all building block of all life and an
oceans. The Redfield Ratio soon important part of many chemical
becomes a benchmark for such processes. Nitrogen is a major
research in all habitats. Individual organisms also constituent of all proteins, while
show differences in phosphorus is crucial for cell
AFTER stoichiometry during their life development and storing energy.
2015 In “Ocean stoichiometry, cycles. Young organisms may An organism’s C:N:P ratio is not
global carbon, and climate,” have different compositions necessarily consistent. Plants have
Robert Sterner highlights from older ones … a variable ratio: they can adjust the
inconsistencies in C:N:P ratios Robert Sterner and balance of their elements according
in phytoplankton, which absorb James J. Elser to their environment. For instance,
more atmospheric carbon in the proportion of carbon in their
low-nutrient, low-latitude ocean chemical makeup can rise on a
surface waters and adjust their particularly sunny day because
ratios accordingly. more photosynthesis occurs—the
process by which they take carbon

See also: Ecophysiology 72–73 ■ The food chain 132–133 ■ Energy flow through
ecosystems 138–139 ■ The foundations of plant ecology 167
The Growth
Rate Hypothesis
Cancer research is one area
Controlling ecological stoichiometry ratios where stoichiometry is now
A locust eats grass that may contain six times being employed. Evidence is
as much carbon as it needs. To get the right growing for a theory called the
balance, it excretes carbon or breathes it out Growth Rate Hypothesis
as CO2. Locusts are widely used in research (GRH), which may help explain
because they are easy to breed. why some cancerous tumors
grow at faster rates than the
rest of the body.
The hypothesis states that
organisms with high C:P
KEY (carbon:phosphorus) ratios,
LOCUSTS GRASS such as fruit flies, have more
Carbon Nitrogen 5:1 33:1
ribosomes in their cells, which
enables them to grow and
dioxide from the air and use the may adjust its digestive enzymes reproduce more rapidly.
sun’s energy to convert it into the and excrete it, store it as fats, or raise Around half of all phosphorus
in an organism is in the form
nutrients they require. its metabolic rate to burn it off,
of ribosomal RNA (rRNA); it is
Higher up the food chain, breathing out the excess carbon as
present in every cell, creating
animals have largely fixed C:N:P CO2. Overuse of such mechanisms proteins to build new cells and
ratios, so they must deploy various to redress a high imbalance can, grow the body. Applying
mechanisms to deal with any however, affect fitness, growth, and biological stoichiometry,
imbalances of chemicals entering reproduction. An animal that eats James Elser and his team
the body. If an insect or animal other animals has less work to do, have shown that fast-growing
herbivore is getting too much carbon because its prey’s C:N:P ratio tumors have a much higher
from its plant diet, for instance, it closely matches its own. However, phosphorus content than
the size of its prey population is still normal body tissue. Such
determined by the plants in its research may help scientists
environment because plants with a understand how tumor growth
high carbon ratio can only support could be controlled.
a small food chain of consumers.

Understanding our world

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

Malignant lung tissue (seen

The desert locust (Schistocerca here) and cancerous colon tissue
gregaria) has to eat vast quantities both had the highest phosphorus
of carbon-rich plants in order to get content in research exploring the
enough nitrogen and phosphorus rapid growth rates of tumors.
to maintain its C:N ratio.



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
in which predators kill and prey are can have an even bigger impact on
Earl Werner (1944–)
eaten. However, American ecologist an ecosystem. Potential prey are
BEFORE Earl Werner and others have shown forced to change their way of life
1966 American ecologist that the mere presence of a predator in order to avoid being killed.
Robert Paine conducts a affects the behavior of prey. In 1990, Werner studied the
series of groundbreaking Apart from apex predators, all effects of green darner dragonfly
field experiments to highlight animals must balance the need to larvae on toad tadpoles. He noticed
the crucial effects of a predator sleep, reproduce, and feed with the that when the predatory larvae
on the community in which
it lives.
1990 Canadian biologists
Steven Lima and Lawrence In the presence of predators …
Dill analyzed the decision-
making of organisms that are
at the greatest risk of being
preyed on by other creatures.
AFTER prey move on to prey spend more time
2008 American behavioral other areas even if there is hiding in sheltered habitats
biologist and ecologist John less food there. than feeding in the open.
Orrock teams up with Earl
Werner and others to produce
mathematical models to
explain the nonconsumptive
effects of predatory animals. Even without preying on
them, predators can cause prey to
fail to thrive.

See also: Evolution by natural selection 24–31 ■ Predator–prey equations 44–49 ■ Ecological niches 50–51
■ Competitive exclusion principle 52–53 ■ Mutualisms 56–59 ■ Optimal foraging theory 66–67

bullfrog’s new behavior gave it

a competitive advantage over the
green frog by making it bigger.

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




Aristotle’s History of A private collection of natural The Natural History

Animals groups living things history curiosities is displayed Museum in London opens its
based on their species, in a at Oxford University’s doors to the public, free of
scala naturae that places Ashmolean Museum, the charge. It now houses 80
organisms into 11 grades. world’s first public museum. million specimens.

C.350 BCE 1683 1881

1665 CE 1758

Micrographia, the richly The 10th edition of Systema

illustrated book by Robert Naturae by Carl Linnaeus
Hooke, reveals microscopic classifies a range of plant
structures to a wider audience. and animal species using
his binomial system.

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,


Norman Myers’s
Carl Woese “biodiversity hotspots”
establishes a new, third concept identifies ten hotspots
category of organisms— where conservation efforts
the prokaryotes. should preserve rare species.

1977 1988

1942 1988 2018

Ernst Mayr develops the Edward O. Wilson coins The IUCN Red List shows that
biological species concept, the term biodiversity more than 26,000 species –
which categorizes species and later identifies the more than 27 percent of all
based on their ability to key human threats those assessed—are at
breed with each other. to biodiversity. risk of extinction.

but it is the 10th edition from 1758 but a population that can breed new domains. As of 2018, about
that founded the modern zoological only among themselves. Mayr went 1.74 million extant plant and animal
naming system. Two volumes of on to explain how if groups within species have been described, but
Linnaeus’ work are devoted to a species become isolated from the estimates of the total number range
plants and animals, which he rest of the population, they may from 2 million to 1 trillion.
divided into classes, orders, genera, start to differ from the rest, and
and species. The binomial system, over time, through genetic drift and The threat to diversity
in which every species is given a natural selection, may even evolve By the late 20th century, however,
generic name followed by a specific into new species. alongside a growing knowledge
name, is still in use today. Linnaeus Modern technological advances, of the scale and critical role of
also wrote a third volume on rocks, including electron microscopy and biodiversity—and of how evolution
minerals, and fossils. mitochondrial DNA analysis, have can destroy species as well as
revealed much information—some create them—American ecologist
Species concepts of it surprising—about the number E.O. Wilson and others made the
Building on Darwin’s theory of of species and the relationships world aware that human activity
evolution by means of natural between them. In 1966, striving to was responsible for causing a rapid
selection, German-American reflect the intricacies of evolution, acceleration in the extinction rate.
evolutionary biologist Ernst Mayr German entomologist Willi Hennig Some have even warned that Earth
cemented the biological concept of proposed a new taxonomic system could be on the verge of a sixth
species in his Systematics and the of clades—groups of organisms mass extinction. Many policies are
Origin of Species (1942). He argued based on a common ancestor. In now being proposed to counter
that a species is not just a group of the 1970s, American biologist Carl this, including the protection
morphologically similar individuals, Woese classified all life into three of biodiversity hotspots. ■



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
organisms according to their uses. of birth. Those in the top grades
Aristotle (c. 384–322 bce)
Egyptian wall paintings from gave birth to live, hot, wet offspring;
BEFORE c. 1500 bce show, for example, that those in the lower grades to cold,
c. 1500 bce Different people understood the medicinal dry eggs. Humans were at the very
properties of plants are properties of many plants. In the top of the scale, with live-bearing
recognized by ancient text History of Animals, written tetrapods (four-legged creatures),
Egyptians. in the 4th century bce, the Greek cetaceans, birds, and egg-laying
philosopher and scholar Aristotle tetrapods lower down. Aristotle
AFTER made the first serious attempt to placed minerals on the bottom
8th–9th centuries ce Islamic classify organisms, studying their grade of his scale, with plants,
scholars of the Umayyad and anatomy, life cycles, and behavior. worms, sponges, larva-bearing
Abbasid dynasties translate insects, and hard-shelled animals
many of Aristotle’s works Features of classification on the levels above.
into Arabic. Aristotle divided living things into
plants and animals. He further
1551–58 Conrad Gessner’s grouped about 500 species of
History of Animals classifies animals according to obvious
the animals of the world into anatomical features, such as
five basic groups. whether they had blood, were
“warm-blooded” or “cold-blooded,” If any person thinks the
1682 John Ray publishes his examination of the rest
whether they had four legs or more,
History of Plants, which lists of the animal kingdom
and whether they gave birth to live
more than 18,000 species. an unworthy task, he must
offspring or laid eggs. He also noted
1735 Carl Linnaeus devises a whether animals lived in the sea, hold in like disesteem
system of binomial names, the on land, or flew in the air. Most the study of man.
first consistent classification of significantly, Aristotle used names Aristotle
organisms, according to which for his groupings that were later
he names every species listed translated into the Latin words
in his Systema Naturae. “genus” and “species”—terms
that are still used by modern
taxonomists to this day.


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

An octopus blends in with its

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

While Aristotle’s system of

classification was rudimentary, it
was based largely on first-hand
observations, many of which were
made on the island of Lesvos. He
recorded things that noone else
had described, including that
young dogfish grew inside their
mothers’ bodies, male river catfish
guard eggs, and octopuses
can change color. Most of his
observations were good—and some
were confirmed only centuries later.

The great chain of being The Swiss doctor Conrad Gessner (reptiles and amphibians); birds;
Despite its limitations, Aristotle’s wrote the first modern register of fish and aquatic animals; and
method of classification heavily animals—also called History of snakes and scorpions. In 1682,
influenced every later attempt at Animals—in the mid-16th century. the English naturalist John Ray
grouping animals and plants until This monumental five-volume work produced the equivalent register
the 18th century. Medieval was based on classical sources but for botany with his History of
Christianity developed his scala included newly discovered species Plants. Within little more than
naturae as a “great chain of being,” from East Asia. It covered the main 50 years, the classification of
with God at the top of a strict animal groups as Gessner saw living things would be completely
hierarchy, humans and animals them: live-bearing quadrupeds transformed by Carl Linnaeus’s
beneath, and plants at the bottom. (mammals); egg-laying quadrupeds Systema Naturae. ■

Aristotle Aristotle was born in Macedonia, scholar Ptolemy and King

ancient Greece. Both his parents Alexander the Great. In 335 bce,
died when he was young, and he he established his own school
was raised by a guardian. Aged at the Lyceum in Athens. After
17 or 18, Aristotle joined Plato’s Alexander’s death in 322 bce,
Academy in Athens, where he Aristotle fled the city, and died
studied for 20 years, writing on on the island of Euboea in the
physics, biology, zoology, politics, same year.
economics, government, poetry,
and music. Later, he traveled Key works
to the island of Lesvos with a
student named Theophrastus 4th century bce
to study the island’s botany and History of Animals
zoology. Much of his History On the Parts of Animals
of Animals was based on On the Generation of Animals
observations he made there. On the Movement of Animals
Aristotle taught both the future On the Progression of Animals



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
astonished. Here, in English in use by the 1660s. The early
Robert Hooke (1635–1703)
scientist Robert Hooke’s seminal instruments were unreliable—due
BEFORE 1665 book, were many detailed to the difficulty of making the
1267 English philosopher illustrations of structures previously lenses—and scientists had to be
Roger Bacon discusses the use hidden from the human eye due to inventive and work around the
of optics for looking at “the their minuscule size. Hooke’s problem. At first, Hooke had
smallest particles of dust” in microscope magnified things by a difficulty seeing his specimens
his Opus Majus Volume V. factor of fifty, but the accuracy of clearly, so he invented an improved
his drawings also owes much to light source, named a “scotoscope.”
1661 Microscopic drawings by his painstaking approach. Hooke Hooke’s book is more than just
English architect Christopher would make numerous sketches an accurate representation of what
Wren impress Charles II, who from many different angles before he saw through the lens; it also
commissions more drawings combining them into a single image. theorizes on what the images reveal
from Robert Hooke. about the workings of the organisms
he studied. For example, when
AFTER looking at a wafer-thin specimen of
1683 Dutch amateur scientist cork, Hooke saw a honeycomb-like
Antonie van Leeuwenhoek pattern, the elements of which he
uses a microscope to observe described as “cells”—a term that is
bacteria and protozoa, and … in every little particle… still used today.
publishes his findings with we now behold almost as
the Royal Society of London. great a variety of Creatures, Microscopic marvels
as we were able before to Micrographia inspired many
1798 Edward Jenner, an reckon up in the whole other scientists to investigate the
English physician and Universe itself. microscopic world. Following
scientist, develops the world’s Robert Hooke notes and diagrams from Hooke’s
first vaccine—for smallpox— book, Dutch scientist Antonie
and publishes An Inquiry into van Leeuwenhoek was able to
the Causes and Effects of the construct his own microscopes.
Variolae Vaccinae. He achieved magnifications of
more than 200 times actual size.


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

Van Leeuwenhoek examined

samples of rainwater and stagnant
pondwater and marveled at the
multitude of life he saw there. He
identified single-celled protozoa,
naming them “animalcules,” and [Micrographia is] … the
went on to discover bacteria. He most ingenious book that I
also made many observations of ever read in my life.
human and animal anatomy, Samuel Pepys
including blood cells and sperm. English diarist
While van Leeuwenhoek
examined water samples, fellow
Dutchman Jan Swammerdam was
placing insects under his own
microscope. He published records
of all manner of insects depicted in
the finest detail and uncovered organ. Grew also spotted pollen
much about their anatomy. grains and noted that they were
Swammerdam’s most influential transported by bees.
work was Life of the Ephemera Since the early days of
(1675), which recorded in great microscopy, devices have grown
detail the life cycle of the mayfly. in sophistication. The electron
In England, Nehemiah Grew microscope, first used in 1931, uses
used microscopy to examine a beams of electrons—rather than
wide range of plants. He was the light—to reveal objects, allowing
first to identify flowers as being scientists an even closer look. The compound eye and brain of a
bee, drawn by Jan Swammerdam and
the sexual organs of plants. In The Electron microscopes provide views published in A Treatise on the History
Anatomy of Plants (1682), Grew of up to one million times actual of Bees, shows the eye exterior (left)
named the stamen as the male size—600 times greater than most and the eye dissected (right), with the
organ and the pistil as the female modern light microscopes. ■ brain cross-sectioned below.

Robert Hooke Born on the Isle of Wight, England, include some early insights into
Hooke showed an early interest the wave theory of light; the
in science. A small inheritance construction of some of the
allowed him to attend the earliest telescopes; and the
prestigious Westminster School, formulation of Hooke’s Law.
where he excelled, earning a place Hooke was also a respected
at Oxford University. There he architect, an activity that made
assisted the natural philosophers him a wealthy man.
John Wilkins and Robert Boyle.
In 1662 Hooke became the first Key works
curator of experiments for the
Royal Society of London. In 1665 1665 Micrographia
he became Professor of Physics at 1674 An Attempt to Prove
Gresham College. the Motion of the Earth
Like many scientists of his 1676 A Description of
day, Hooke had a broad range of Helioscopes and Some
interests. His achievements Other Instruments




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
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
Plantae, and Animalia. the names of things, cause discrepancies in
the knowledge of scientific knowledge.
them is lost.


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

been known by long impractical

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

Key works

1735 Systema Naturae

1737 Flora Lapponica
1751 Philosophia Botanica
1753 Species Plantarum



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.
1976 The Selfish Gene by

Richard Dawkins popularizes y the early 20th century, it some point become separated by
gene-centered evolution: natural was accepted that multiple geography, mate choice, feeding
selection at a genetic level. species could evolve from a strategies, or other means, and then
1995 The Beak of the Finch common ancestor. However, it was begin to change through natural
not clear how this evolution process selection or genetic drift. Over time,
by Jonathan Weiner follows
actually occurred. In fact, there was as a result of this initial separation,
the work of biologists Peter
some debate about precisely what a two distinct species evolve, which
and Rosemary Grant on the
“species” was. In 1942, evolutionary cannot interbreed. This type
Galapagos Islands. biologist Ernst Mayr proposed a of speciation commonly occurs
2007 Massimo Pigliucci and new definition of species: groups in small populations of creatures
Gerd B. Müller use the term of interbreeding natural populations on remote islands.
“eco-evo-devo” to suggest how that are “reproductively isolated
ecology is among the factors from other such groups.” Key differences
affecting evolution. What this means is that two The biological species concept is
populations of the same species primarily focused on the breeding
living in the same area may at potential between organisms. Two


See also: Evolution by natural selection 24–31 ■ The role of DNA 34–37 ■ The selfish gene 38–39
■ Competitive exclusion principle 52–53

organisms may appear identical Sometimes, too, different animal

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

Alternative species concepts

Although Mayr’s idea about be based on genetics, such as
biological speciation is perhaps DNA or RNA base sequences, or
the most common way to define on phenotypes, such as the size
species and explain how they of certain body parts or
evolve, it is far from the only particular markings, such as the
one. In fact, there are more than arrangments of spots on insects’
20 recognized species concepts, wings. The evolutionary species
ranging across two broad groups: concept is based on species
typological and evolutionary lineages. A species is defined as
concepts. Typological species the organisms that share a
Male fireflies are an example of concepts are based on the idea lineage from the time when the
a typological species. They emit a that a population of individuals of species initially split off until
pattern of flashes to attract females, the same type—or sharing the extinction, or until an additional
who recognize their species’ code same set of traits—are what splitting off and creation of a
and flash back—if they wish to mate. makes up a species. The traits can new species.



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
microscopic structure of living complex cells).
Carl Woese (1928–2012)
things, biological diversity was In the 1970s, the American
BEFORE split simply into animal-like and biologist Carl Woese claimed that
1758 Systema Naturae (10th plant-like organisms. Then, in the even this system failed to account
edition) by Carl Linnaeus 20th century, better microscopes for the diversity among microbes—
classifies known life into two began to reveal deeper differences the smallest living things. He
kingdoms: animals and plants. that could not be seen with the focused on ribosomes—minuscule
naked eye. By the 1960s, picking
1937 French biologist Edouard up on an idea first proposed by
Sulfur-dependent archaea
Chatton divides life into Edouard Chatton in the 1930s, the organisms thrive in the hot geothermal
prokaryotes (bacteria) and need for a new division of living pools of Yellowstone National Park,
eukaryotes (organisms with things emerged, placed between Wyoming, in conditions that would
complex cells). prokaryotes (such as bacteria, kill most other organisms.

1966 German biologist Willi

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


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


Edward O. Wilson (1929–)

1993 The UN proclaims
December 29 as the
International Day for


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


David Quammen explores
the nature of evolution and
extinction as habitats become
more and more fragmented.


2014 The Sixth Extinction
by environmental journalist
Elizabeth Kolbert shows how
humans are causing a sixth
mass extinction of species.
2016 In Half-Earth, Edward
Wilson proposes that Earth
can be saved by dedicating
half of it to nature.

iodiversity is the variety of
life on Earth—in all forms
and at every level, from
genes to microbes to humans and
all other species, including those
yet to be discovered. Humans rely
on biodiversity for food and fuel,
shelter, medicine, beauty, and
pleasure. For other species, it also
provides nutrients, seed dispersal,
pollination, and reproductive
success. No living thing could
survive without biodiversity.
Ecologists have identified
growing threats to biodiversity,
many of them driven by human
actions. The current rate of species
extinction is thought to be up to
1,000 times greater than it was


See also: Biodiversity hotspots 96–97 ■ Animal ecology 106–113 ■ Island biogeography 144–149 ■ Biodiversity and
ecosystem function 156–157 ■ Biomes 206–209 ■ Mass extinctions 218–223 ■ Deforestation 254–259 ■ Overfishing 266–269

The effects of human activity on biodiversity animals because they may no

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


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


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

against natural disasters and

human-made shocks, including
climate change, and provide
recreational, medicinal, and
biological resources.
Although the threats to
biodiversity from human activity
are serious, ways to protect it
are being developed. Foremost
is a “sustainable” approach to
harvesting and agriculture that
allows species—such as fish, trees,
When the rate of harvest exceeds pollution and climate change. More or crops—to be maintained at a
the rate of replenishment through than 80 per cent of the species on stable level and even increased
either reproduction or human the IUCN’s Red List are affected by over time. Official protected status
activities such as tree planting, more than one of the five major for areas of land, water, and ice
the harvest is not sustainable, biodiversity threats. can help sustain threatened
and without regulation could result Biodiversity maintains the species, while national and
in the extinction or extirpation health of the ecosystems of the international agreements and
of species. planet. Ecosystems are a delicate negotiations can mitigate the
A study of the IUCN’s Red List balance of living creatures, both impact of both legal and illegal
in 2016 showed that 72 per cent plant and animal, as well as the trade, such as poaching. Public
of species listed as threatened or soil, air, and water in which they education also helps people to
near-threatened are harvested at live. Healthy ecosystems provide better understand their potential
a rate that means their numbers resources that sustain human and impacts on biodiversity and how
cannot be balanced by natural all other life, improve resilience to protect it for future generations. ■
reproduction or regrowth. Some
62 per cent of species are at risk Anthropogenic biomes grouped into six main
from agricultural activity alone, categories: dense settlements;
such as livestock farming, tree The biosphere—all the areas of villages; croplands; rangeland;
felling, and the production of Earth and its atmosphere that forested; and wildlands.
crops for food, fuel, fibres, and contain living things—consists Unlike other biomes, which
animal fodder. of biomes, which are large can range across continents,
ecosystems based on a specific anthropogenic biomes are a
Protecting biodiversity environment, such as desert or mosaic of pockets over Earth’s
In reality, the five HIPPO threats tropical rainforest. The impact surface. According to ecologists,
identified by Wilson are interrelated, of human actions on biodiversity more than 75 per cent of Earth’s
and there is generally no single and the consequent reshaping ice-free land has been affected
of much of the planet have led by at least some form of human
reason why any particular species
ecologists to reassess biomes activity, particularly in dense
is endangered. Agricultural
and suggest that a designation settlements (urban areas), which
development, for example, can of anthropogenic (manmade) account for over half the world’s
not only destroy a habitat, but can biomes is now necessary. population, and villages (dense
also releases greenhouse gases into Anthropogenic biomes are agricultural settlements).
the atmosphere, contributing to air



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,
animal and plant species. The term Myers argued that priorities had
Norman Myers (1934–)
was coined in 1988 by Norman to be set to establish where to
BEFORE Myers, a British conservationist, concentrate resources to conserve
1950 Theodosius Dobzhansky to describe areas that are both as many lifeforms as possible.
studies plant diversity in biologically rich and deeply
the tropics. threatened. Facing the huge Defining hotspots
Initially, Myers identified ten
AFTER hotspots crucial for conserving
2000 Myers and collaborators The lush hillsides and forests of
Arunachal Pradesh, India, are part plant species that were endemic
reevaluate the list of hotspots of the Indo-Burma biodiversity hotspot. (did not grow anywhere else on
and add several new ones, The area contains some 40 per cent Earth). By 2000, he had refined the
bringing the total to 25. of India’s animal and plant species. concept to focus attention on
2003 An article in American
Scientist criticizes the
concentration of conservation
effort on hotspots, saying that
this neglects less species-rich
but still important “coldspots”.
2011 A team of researchers
confirm the forests of east
Australia as the 35th hotspot.
2016 The North American
coastal plain is recognized
as meeting the criteria for a
global biodiversity hotspot—
and becomes the 36th.


See also: Human activity and biodiversity 92–95 ■ The ecosystem 134–137
■ Deforestation 254–259 ■ Sustainable Biosphere Initiative 322–323

looks like an antelope; it was seen

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




Louis Pasteur reveals Charles Elton publishes

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

1590 1866 1927

1676 1885

Antonie van Albert Frank coins the

Leeuwenhoek term “mycorrhizae,” in
identifies “animalcules,” reference to the symbiotic
opening up the field relationship between
of microbiology. fungi and tree roots.

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.


In Man Meets Insect Thermoregulation In the US, the Human

Dog, Konrad Lorenz by Bernd Heinrich Microbiome Project is
describes the instinctive explains how insects launched, to map all the
behavior of animals and are able to control their microbes associated with
its evolutionary origins. own temperature. a healthy human body.

1949 1981 2007

1947 1960 2005

David Lack publishes Jane Goodall sets up The first gene mapping
an article on variations a camp in Tanzania to confirms that humans
in bird clutch size as research chimpanzees in share 97–99 percent of
an evolutionary the wild, discovering many their DNA with the other
adaptation. traits shared by humans. great apes.

Ethology, which looks at animal Field observations are a key tool in a red spot on a parent’s beak when
behavior and its evolutionary basis ethological research. In the 1940s, they want food, will tap colored
and development, is a major British ornithologist David Lack marks painted on a model beak.
component in the modern study investigated the factors controlling
of organisms. Back in 1837, British the number of eggs birds laid Human traits
entomologist George Newport (clutch size). His food limitation As well as these short-term studies,
discovered that moths and bees hypothesis states that the number British primatologist and ethologist
could raise the temperature of their of eggs laid by a species has Jane Goodall conducted field
thorax by quivering their muscles. evolved to match the food available. observations over a longer period,
From the 1970s onward, German- Evolutionary pressure has created studying chimpanzees in Tanzania
American entomologist Bernd a correlation between clutch size from 1960 to 1975. Her findings
Heinrich and others uncovered and food availability. challenged the view that human
more thermoregulatory adaptations Austrian zoologist Konrad behavior is totally unique in the
that have helped insects thrive. Lorenz and Dutch biologist Nikolaas animal world, and indicated that
As heterotherms, they are able Tinbergen also studied animals in chimps are behaviorally closer
to maintain different temperatures the wild to help understand their to people than had generally been
in different parts of the body. behavior. Lorenz’s 1949 work Man assumed. She noted, for example,
Modern research now combines Meets Dog explains the loyalty that chimps display a whole range
laboratory experiments, field of a pet dog to its owner in terms of facial expressions and other body
observation, and new technology of canines’ instinctive loyalty language to indicate their mood, are
such as infrared thermography to their pack leader in the wild. toolmakers and users, often behave
to understand insect behavior Tinbergen’s field experiments cooperatively, and sometimes go
in ever more detail. showed how gull chicks, which tap into battle against rival groups. ■



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
in every environment, living in soil, 17th century, using the recently
Louis Pasteur (1822–95)
water, and air. Some microbes invented microscope. These
BEFORE cause disease but most are vital for studies revealed a previously
1683 Dutch amateur scientist life on Earth. Among other things, unknown world teeming with
Antonie van Leeuwenhoek they break down organic matter so microbiotic life. Around the same
uses a microscope to observe that it can be recycled back into time, the word “germ,” originally
bacteria and protozoa. the ecosystem. meaning “seed,” was first used
Trillions of microbes also live on to describe these tiny organisms.
1796 Edward Jenner carries and in the human body. The most
out the first vaccination, using common of these microbes are Fighting disease
the cowpox virus to protect beneficial bacteria, which aid the Some 17th and 18th-century
against smallpox. digestion of food, produce vitamins, scientists believed that certain
and help the immune system find “germs” might cause diseases, but
AFTER and attack more harmful microbes. the prevailing view was that such
1926 American microbiologist maladies were the spontaneous
Thomas Rivers distinguishes result of inherent weakness in an
between viruses and bacteria. organism. It was not until the
1928 While studying painstaking laboratory work of the
influenza, Scottish 19th-century French chemist Louis
Pasteur that the “germ theory of
bacteriologist Alexander
Microbes are the worker bees disease” was proved.
Fleming discovers penicillin.
that perform most of the Pasteur began by looking at the
2007 An inventory of all important functions alcohol fermentation process. He
the microbes associated discovered that sourness in wine
in your body.
with a healthy human body was caused by external agents—
Dr Robynne Chutkan, microbes, or germs. A crisis in the
is completed. Microbiome expert and author
French silk industry, caused by an
epidemic among silkworms, then
allowed Pasteur to isolate and
identify the microorganisms that
caused the particular disease.


See also: Classification of living things 82–83 ■ The microbiological
environment 84–85 ■ The ecosystem 134–137

animal or human, was particularly

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

1870 Studies on Silk Worm

1878 Microbes: Their Role in
Fermentation, Putrefaction,
and the Contagion
1886 Treatment of Rabies




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


See also: Evolution by natural selection 24–31 ■ Mutualisms 56–59
■ The ecosystem 134–137 ■ Energy flow through ecosystems 138–139
Mycorrhizae as
pollution indicators
Mycorrhizal fungi are not only
Beneficial exchange between good for the health of plants—
mycorrhizae and plant roots they can also act as indicators
of the health of the entire
Mycorrhiza Plant environment. Laboratory
Supplies sugar from
photosynthesis experiments with these fungi
have shown that some grow
badly in the presence of toxins,
Connects plants in an which means that they can be
extensive network used to detect pollutants in
the air or soil. For instance,
some fungi fail to grow when
Increases uptake of exposed to heavy metals such
The mutualistic as lead or cadmium, and
relationship between water and nutrients
because different kinds of
mycorrhizae and plants fungi react differently to
is highly evolved. As
Allows plants to share environmental change, certain
many as 90 percent of
all plant species rely on nutrients with others species can be used to identify
fungi for nutrients and specific kinds of pollution.
protection. In return, Mycorrhizae are also useful
plants supply the fungi Boosts protection indicators of the health of their
with a vital food source. against soil diseases native habitat. Many form
cauliflower-like growths
on tree roots, but these are
a much greater amount of minerals. fungi in this way. Trees supported smaller in polluted soil. The
When the fungal hyphae attach to by mycorrhizae are more resistant trees themselves may also
the plant roots, they extend the root to drought and disease, and can respond to pollution with
system, causing extra nutrients to even communicate alarm signals by weaker shoot growth, but
seep into the plant. releasing chemicals in response to the mycorrhizal response is
Albert Frank realized that this attack by herbivores. This fungal more acute and serves as a
partnership worked both ways. It network connecting trees has been valuable early-warning sign
was a winning combination for both dubbed “the wood-wide web.” ■ of a habitat in decline.
plant and fungus. In exchange for
passing on a share of its minerals,
the fungus receives sugar from the
plant—made by photosynthesis in
the leaves and transported to the
roots via the plant’s sap. This boosts
the nutrient supply that the fungus [the fungus] performs
derives from dead organic matter. a “wet nurse” function
and performs the
Ancient networks entire nourishment
Fossils of plants dating from 400 of the tree from
million years ago—when vegetation the soil.
was first spreading across dry Albert Frank
land—show traces of fungal threads. Weak growth in the russet
This suggests that the mycorrhizal brittlegill, a mycorrhizal fungus
of European and North American
partnership was key to the evolution spruce forests, can be an early
of terrestrial life. Today, the majority indicator of habitat air pollution.
of plant species continue to rely on



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
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
1960 American ecologist and
philosopher Garrett Hardin
publishes an essay in the
magazine Science in which
he states that “every instance Whale
of apparent coexistence must Albatross

be accounted for.” Krill

1973 Australian ecologist
Robert May publishes Stability
and Complexity in Model Arrow worm Leopard
Ecosystems, in which he uses seal
mathematical modeling to Blue-green Radiolarians
demonstrate that complex bacteria

ecosystems do not necessarily Penguin

lead to stability. Schooling fish


A food web is a graphic depiction

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


See also: Keystone species 60–65 ■ The food chain 132–133 ■ The ecosystem
134–137 ■ Energy flow through ecosystems 138–139 ■ Trophic cascades 140–143
Forecasting the effects
of climate change

in nature, so if the population Ecologists examine changes

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


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


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

Niches and competition

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

A true specialist, a koala bear

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


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

Ecological pyramids represent Microscopic organisms, including

quantifiable data in an ecosystem. these diatoms, form a significant part
Numbers show the population size of of all ecological pyramids. Their huge
individual species in a trophic level; numbers and rapid reproduction
biomass, their relative presence; and provide mass and energy for the
energy, who eats what and how much. species higher up the pyramid.


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




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.
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.
1948 David Lack extends his
theory of optimal clutch size
in birds to include litter size
in mammals.
1954 Lack develops his food
limitation hypothesis further
in The Natural Regulation
of Animal Numbers, to
encompass birds, mammals,
and some insect species.
1982 Tore Slagsvold proposes
the nest predation hypothesis,
which states that clutch size
is related to the likelihood of
the nest being attacked.


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

■ Ecological resilience 150–151

Siblicide and the
Blue-footed Booby

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

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




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


See also: The selfish gene 38–39 ■ Field experiments 54–55 ■ Keystone species 60–65 ■ Animal ecology 106–113 ■ Clutch
control 114–115 ■ Using animal models to understand human behavior 118–125 ■ Thermoregulation in insects 126–127

pattern” (FAP). A FAP has set Four elements of ethological experimentation

characteristics. It is species-
specific; it is repeated in the same
way every time and is not affected
by experience. The triggers for the Development
behavior (“sign stimuli”) are highly What stage is the animal
What triggered
specific and may involve a color, in its life cycle, and does the
the behavior in
pattern, or sound. For example, male behavior change as
the first place?
sticklebacks respond aggressively the animal develops?
when another male enters their
streambed patch. Ethologists
suggest this is triggered by seeing
the male’s red underbelly.
Nikolaas Tinbergen found When studying
that some an artificial sign stimuli animal behavior,
work better than the real thing. He scientists consider
investigated the begging behavior
these four elements
of herring gull chicks, which peck
at a red spot on the parent gull’s
beak to make it regurgitate food.
He found that chicks will also peck
at a model of the gull’s beak, yet
when they were offered a narrow Function
Evolution How does the behavior
red pencil with three white lines How is the behavior
at the end, the chicks pecked at increase the animal’s
related to the animal’s chance of survival or
this even more enthusiastically. evolution or ancestry?
Tinbergen called this a reproductive success?
“supernormal stimulus,” showing
that instinctive animal behavior
can be manipulated artificially. ■

Konrad Lorenz Born in Vienna, Austria, Lorenz and other birds, as well as
was enthralled by animals from mammals, is instinctive and
an early age and kept fish, birds, occurs shortly after birth. Lorenz
cats, and dogs. The son of an demonstrated the theory by
orthopedic surgeon, he studied quacking like a duck at newly
medicine at Vienna University, hatched ducklings. He soon
graduating in 1928, and gained had a flock of ducklings that
his Ph.D. in zoology in 1933. His followed him everywhere.
numerous pets became the first
subjects of his studies. Lorenz is Key works
perhaps best known for describing
the phenomenon known as 1952 King Solomon’s Ring:
“imprinting.” This is when a New Light on Animal Ways
newly hatched chick bonds with 1949 Man Meets Dog
the first thing it sees (usually its 1963 On Aggression
parent) and will follow it around. 1981 The Foundations for
The behavior, seen in ducks Ethology



The Primates Tree
Humans and their
KEY FIGURE closest relatives,
Jane Goodall (1934–) chimpanzees, are both
primates. This shows
BEFORE 750,000–550,000 ya how primates have
1758 Carl Linnaeus, the father evolved over the last
66 million years.
of taxonomy, dares to classify 6 mya
humans within the rest of
nature, calling us Homo
sapiens (“wise man”).
1859 Charles Darwin’s apes
theory of evolution further
challenges the established Lesser
view that man is different
from the animal kingdom. Old world
1963 Konrad Lorenz publishes New world
On Aggression, proposing 35 mya
that warlike behavior in
humans is innate. Primates
40 mya
1967 Desmond Morris,
a British zoologist and
ethologist, publishes The 65 mya
Naked Ape: A Zoologist’s
Study of the Human Animal,
a major study that describes Modern Chimpanzee/
human Neanderthal Gorilla Bonobo Orangutan
human behavior in the
context of the animal kingdom.
Gibbon Monkey Baboon Tarsier Lemur Loris

odern molecular studies scientific community remained
mapping the genomes convinced that humankind was
of humans and other different from the rest of nature.
animals have confirmed a theory It was largely the work of British
that was first suggested by Charles primatologist Jane Goodall that
Darwin in the mid-19th century— opened our eyes to the similarities
that we share a common ancestor between chimps and man. In 1961,
with the great apes. Today, few in an excited communication
In reality, we are Pan narrans, scientists would dispute that the to her mentor, Louis Leakey,
the storytelling chimpanzee. common chimpanzee (Pan Goodall announced an observation
Terry Pratchett troglodytes) and the bonobo or that would shake the scientific
British fantasy author pygmy chimpanzee (Pan paniscus) establishment: she had seen
are our closest living relatives. The a chimp using a tool. It was the
study of these animals therefore first time this behavior had
offers us a unique chance to learn been documented and it would
about ourselves and the origins of challenge perceived ideas of what
our behavior. Yet for many years the it means to be human.


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

Goodall’s knowledge of natural

history had impressed Leakey
on their first meeting in 1957 and
he offered her a job studying the
behavior of chimpanzees. As an
anthropologist and paleontologist,
Leakey believed in evolutionary
theory, which proposed that humans
and the great apes—chimpanzees,
bonobos, gorillas, and orangutans—
in the family Hominidae (Great
Apes), share a common ancestor.

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

Jane Goodall Born in London in 1934, Jane transformed our understanding

Goodall’s first meeting with a of chimpanzees and challenged
chimp was a stuffed animal that perceived ideas of our own place
her father named Jubilee. She was in the natural world. In 1965, she
interested in animal behavior from earned a Ph.D. in ethology from
an early age—once, she hid in a Cambridge University. Her many
henhouse for hours so that awards include France’s Legion
she could watch a chicken lay of Honor, given to her in 2006.
an egg. She left school at 18 and
worked in various jobs, before Key works
going to Kenya in 1957 and
meeting paleoanthropologist 1969 My Friends the Wild
Louis Leakey. With his support, Chimpanzees
in 1960 Goodall set up a research 1986 The Chimpanzees of
base in Gombe, Tanzania, where Gombe: Patterns of Behavior
she was to study chimpanzees 2009 Hope for Animals and
until 1975. Her work radically Their World


camp, the chimps fled from her, and then using them in termite Jane Goodall working, notebook in
but they then began to forget she mounds; the chimps were not only hand, at Gombe National Park in 2006.
was there. using tools but making them. The pioneering primatologist continues
her lifelong commitment to protect
Goodall sat for many hours endangered chimpanzees.
observing the chimps, keeping her Chimp technology
distance and quietly making field Goodall went on to witness nine
notes. One morning in November different tools being used by instead of numbers, suggesting
1961, she noticed a chimpanzee she chimps in the Gombe community. that her fieldwork was less than
called David Greybeard sitting over At the time, scientists questioned rigorous. Since then, however, many
a termite mound. He was poking Goodall’s methods and ridiculed other studies around the world have
blades of grass into the mound, her for giving the chimps names corroborated her findings: chimps in
pulling them out, and then putting the Congo have been observed
them into his mouth. She watched stripping twigs to use in termite
for some time before the chimp mounds; chimps in Gabon have
moved off. On reaching the spot been seen heading into the forest
where the chimp had been sitting, with a five-piece “toolkit” that
Goodall saw discarded grass stems included a heavy stick for opening
lying on the ground. Picking one up I viewed my bee hives and pieces of bark for
and poking it into the mound, she fellow man not scooping up the honey. In Senegal,
found that the agitated termites bit as a fallen angel, hunting parties of chimps have been
onto the stem. She realized the but as a risen ape. observed traveling with sticks that
chimp had been “fishing” for Desmond Morris they chew to a sharp point and use
termites with the grass stems, and British Zoologist like spears to kill bush babies.
transferring them into his mouth.
From talks with Leakey, Goodall More alike than different
knew this was a major discovery. Ethologists take behaviors studied
She also saw chimps modifying thin across several species to formulate
twigs by stripping them of leaves generalizations that apply to many


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

An alpha male’s body language

says “keep away” to begging chimps
wanting a share of his prey in Gombe
National Park. The main source of prey
for chimps is the colobus monkey.


Conservation of chimpanzees
According to the Jane Goodall chimp communities when roads
Institute in Tanzania, the number are built through their territories.
of chimpanzees living in the wild Roads also encourage another
has plummeted over the last damaging activity—hunting
century. In 1900, there were an for bushmeat, a highly valued
estimated 1 million chimpanzees meat in Africa that includes
in Africa; today, there are fewer great apes. Roads enable
than 300,000. Habitat loss due to hunters from towns to travel
a rising human population in need directly into the bush. The
Orphaned chimps—their mothers of more space has had a huge protection of chimps focuses
killed for bushmeat—walk along a impact, as have industries such on land conservation and on
mud track with their keeper at a as logging and mining, which raising awareness both locally
conservation center in West Africa. destroy habitat and fragment and across the globe.

Such behavior has implications Although chimps sometimes hunt for the war was unclear; some
for human evolution. Science has alone, hunting tends to be a group researchers blamed the feeding
long questioned why and when activity. Chimps rampage through stations Goodall had set up in the
humans first began eating meat. the forest, coordinating their area, which may have encouraged
From prehistoric stone tools and positions and surrounding their unnatural congregations of chimps.
marks on bones, paleontologists prey. After the hunt, the food is The answer to the mystery came
know that the early hominids shared. This shows how early in March 2018, when a research
were using stone tools to cut meat ancestors of humans may have team at Duke and Arizona State
from animals bones 2.5 million developed cooperative behavior, Universities, US, digitized Goodall’s
years ago, but it is not known a factor that may have contributed meticulous check sheets and field
what they were eating between to their evolutionary success. notes from 1967 to 1972 and fed
then and 7 million years ago, them into a computer in order to
when the common ancestor Chimp warfare analyze the social networks and
of chimpanzees and humans A shocking revelation that came alliances of all the male chimps.
is thought to have lived. out of the Gombe camp was that Their findings revealed that the
It is likely that these early chimps are capable of violence, fracture in the community occurred
hominids hunted prey. Although murder, and in particular warfare— two years before the war broke
they did not have large canine once believed to be the preserve out, when an alpha male Goodall
teeth like chimpanzees, these are of humans. Between 1974 and 1978, called Humphrey took over the
not necessary for hunting and Jane Goodall watched as her troupe, alienating two other high-
killing small prey. Biologists have peaceful community of chimps
observed that chimps hunting fractured into two rival groups that
colobus monkeys grab them from then waged savage war on each
the trees and then kill them by other. Goodall was deeply upset
repeatedly thumping the bodies about the chimps’ activity, which
on the ground; early hominids included ambushes, kidnappings,
could have hunted and killed and bloody murder. The trigger
in a similar fashion long before
the earliest known tools.
Chimps may fight over territory
in order to acquire more resources
Cooperative behavior or mates, but some primatologists
Another aspect of chimps’ hunting maintains that such aggression is
behavior that is similar to that unnatural and provoked by human
of humans is the social element. impact on their habitat.


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

Fights over resources

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




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


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




Nelson Hairston,
Richard Bradley Charles Elton Frederick Smith, and
describes how plants, develops the idea of The journal Ecology Lawrence Slobodkin’s
pollinating insects, the food web in Animal posthumously publishes “green world hypothesis”
and insectivores rely Ecology and introduces Raymond Lindeman’s article argues that the predator–
on one another in a the concept of the “The trophic-dynamic prey balance is key to
food chain. ecological niche. aspect of ecology.” flourishing ecosystems.

1718 1927 1942 1960

1859 1935 1957

Charles Darwin Arthur G. Tansley coins the G. Evelyn Hutchinson

describes food term ecosystem, arguing establishes the concept
webs in his On the that an environment and all of niche breadth at
Origin of Species. its living organisms have the Cold Spring
to be seen as a single, Harbor Symposia on
interactive whole. Quantitative Biology.

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


Hal Caswell proposes Scientists at the

a “neutral” theory Biodiversity and
John Maynard Smith of biodiversity, suggesting Ecosystem Function
defines his that competitors are often conference in Paris
Evolutionarily Stable equal, and chance plays examine how the
Strategy (ESS) theory the deciding role in loss of species
in On Evolution. what does or doesn’t thrive. impacts ecosystems.

1972 1976 2000

1967 1973 1980 2015

Robert MacArthur Crawford Stanley Robert Paine coins the term A study of grassland plants
examines the Holling uses the “trophic cascade” after his suggests that biodiversity
biodiversity of term ecological field experiments show the increases an ecosystem’s
isolated communities resilience to show how effect on an ecosystem resistance during
in The Theory of ecological systems when a keystone species and resilience after
Island Biogeography. survive change. is removed. climate events.

surrounded by ocean but also evolutionarily stable strategy (ESS) after a trauma. Ecologists now
“islands” of distinct habitat to describe the best behavioral understand that ecosystems can
surrounded by a very different strategy for an animal competing have more than one stable state,
environment—is so important with others living in its vicinity. and that resilient systems are not
in ecology. In the US in the 1960s, This strategy depends on how the always good for biodiversity.
Edward O. Wilson and Robert other animals behave and is When the populations of many
MacArthur discovered key factors determined by the animal’s genetic species are declining or becoming
determining species diversity, success—if it makes the wrong locally extinct, ecologists are once
immigration, and extinction on decisions, it will not live long and more focusing their attention on
islands. James Brown later did cannot pass on its genes. The ecosystem resilience. Many,
similar work on animal populations overall balance between the including French ecologist Michel
in isolated patches of forest ridge in evolutionarily stable strategies of Loreau, believe that if diversity in
California. Such work has showed all the animals in an ecosystem is an ecosystem is reduced, the whole
how to identify species most at risk called the evolutionarily stable state. system will be less likely to resist
of extinction due to isolation. Canadian ecologist Crawford major impacts such as the effects of
Stanley Holling introduced the idea climate change. Today, Loreau and
Stability and resilience of resilience—how ecosystems others are working toward finding a
One major contribution to the persist in the wake of disruptive new general theory that can
understanding of ecosystem changes such as fire, flood, or explain the relationship between
dynamics was the concept of the deforestation. A system’s resilience ecosystem biodiversity and
evolutionarily stable state. In the can be seen in its capacity to absorb resilience in order to understand
1970s, British biologist John disturbance, or the time it takes to and combat the effects of today’s
Maynard Smith used the term return to a state of equilibrium environmental challenges. ■




The food chain

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

See also: Predator–prey equations 44–49 ■ Mutualisms 56–59 ■ Keystone species 60–65 ■ Optimal foraging theory 66–67
■ Animal ecology 106–113 ■ The ecosystem 134–137 ■ Trophic cascades 140–143 ■ Ecological resilience 150–151

Each species has a

specific place in nature,
in geographic location
and in the food chain.
Carl Linnaeus

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



Arthur Tansley

1864 George Perkins Marsh,

SOURCES OF an American conservationist,

publishes Man and Nature,


which hints at the concept
of ecosystems.
1875 Austrian geologist

Eduard Suess proposes
the term “biosphere.”


1953 American ecologists
Howard and Eugene Odum
develop a “systems approach”
to studying the flow of energy
through ecosystems.
1956 American ecologist
Paul Sears highlights
the role of ecosystems
in recycling nutrients.
1970 Paul Ehrlich and Rosa
Weigert warn of potentially
destructive human
interference in ecosystems.

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.

See also: Animal ecology 106–113 ■ The food chain 132–133 ■ Energy flow through ecosystems 138–139 ■ The biosphere
204–205 ■ The Gaia hypothesis 214–217 ■ Environmental feedback loops 224–225 ■ Ecosystem services 328–329

Tropical coral reefs are some of the

most diverse ecosystems of all, full of
fish, sea turtles, crustaceans, mollusks,
and sponges, as well as corals.

This idea had been developing

long before Arthur Tansley
published his influential paper
on the subject in 1935. As early
as 1864, conservationist George
Perkins Marsh, in his book Man
and Nature, had identified “the
woods,” “the waters,” and “the
sands” as different types of habitat.
He examined how the relationship
between them and the animals and
plants that lived in them could be
upset by human activity.

Interconnected systems
By the 20th century, the idea had
taken hold that these and other
environments could be understood a single unit, and using the term nitrogen, and soil minerals, which
as discrete entities, with distinctive “biome” to describe the whole are essential to the functioning of
interactions between the living complex of organisms inhabiting the systems as a whole. The biotic
and inert elements within them. In a given region. components within an ecosystem
1916, American ecologist Frederic Tansley envisaged ecosystems not only interact with one another,
Clements built on this idea in his as being made up of biotic (living) but also with the abiotic parts.
work on plant succession, referring elements and abiotic (nonliving) Thus, within any given ecosystem,
to a “community” of vegetation as elements such as energy, water, the organisms adapt to both the ❯❯

Arthur G. Tansley A free-thinking Fabian socialist He retired in 1937, but

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


biological and physical elements matter: plants absorb carbon
of the environment. The different dioxide (CO2) from the atmosphere
types of ecosystem can be defined and nutrients from the soil to grow.
by their physical environments. These plants release life-sustaining
There are four categories of oxygen into the atmosphere by
ecosystem: terrestrial, freshwater, respiration, and provide food for
marine, and atmospheric. These animals. Animal excreta and dead
can be further subdivided into matter also release carbon, and
various types according to different provide material to be decomposed
physical environments and the by bacteria and fungi, in turn
biodiversity within them. Terrestrial providing soil nutrients for plants.
ecosystems, for example, can be Arthur Tansley also argued
subdivided into deserts, forests, that these internal processes
A small glacial lake, or tarn, in the
grasslands, taigas, and tundras. within an ecosystem conform to English Lake District. Each tarn has
what he described as “the great an ecosystem that varies according
Dynamic feedback universal law of equilibrium.” to many factors, including the degree
Tansley’s most important insight Self-regulating, these processes of nutrient enrichment in the water.
was that these discrete communities have a natural tendency toward
of living and nonliving components stability. The cycles within an Each ecosystem is located in a
form dynamic systems. In a ecosystem contain feedback loops particular area, with characteristics
terrestrial ecosystem, for example, that correct any fluctuations from unique to its environment, and
the organisms interact to recycle a state of equilibrium. behaves as a self-contained and
self-regulating system. Together,
the patchwork of ecosystems
The dynamic transfer of energy across the globe form what
Austrian scientist Eduard Suess
In this ecosystem, plants use the Sun’s energy
for photosynthesis. As shown by the pale arrows,
called the biosphere—the sum
this energy is passed on—to herbivores, who eat total of all ecosystems.
plants; to predators, who eat herbivores; and to
saphrophytes, who take energy from decomposing External factors
remains and transfer nutrients to the soil. Various external factors, such as
At each stage, some energy is lost as heat. climate and the geological makeup
Heat of the surrounding environment,
can affect an ecosystem. One
Heat constant external force that affects
Plants all ecosystems is the Sun. The
(producers) Heat supply of energy that it provides
enables photosynthesis and the
capture of CO2 from the atmosphere;
Frog some of this energy is distributed
(predator) through the ecosystem and through
the food chain. In the process, some
of this energy dissipates as heat.
Other external factors, however,
(herbivores) can arise unexpectedly to create
Saphrophytes pressures on ecosystems. All
and parasites ecosystems are subjected to
external disturbances from time
to time, and must then go through
a process of recovery. These
disturbances include storms,

more fragile, and if they are wholesale destruction of habitat
disturbed may never be able and the consequent depletion of its
to regain their equilibrium. biodiversity. In addition, some have
The resistance and resilience suggested that human influence
of an ecosystem is generally has created a new category
There is no waste in thought to be related to its of ecological systems, dubbed
functioning … ecosystems. biodiversity. If, for example, “techno-ecosystems.” For example,
All organisms, dead or alive, there is only one species of plant “cooling ponds” are manmade
are potential sources of food performing a particular function ponds, built to cool down nuclear
for other organisms. in the system, and that species is power plants, but they have become
G. Tyler Miller not frost-resistant, an abnormally ecosystems for aquatic organisms.
Science writer severe winter could deplete the The relationship between
species enough to have a major humans and natural ecosystems
impact on the system as a whole. is not all negative. In recent years,
In contrast, if there are several scientific data has fueled public
species with that role in the awareness of the benefits that
system, it is more likely that one ecosystems afford humankind,
will be resistant to the disturbance. including the provision of food,
earthquakes, floods, droughts, and water, nutrients, and clean air,
other natural phenomena, but are The human factor as well as the management of
increasingly the result of human Some disturbances can be severe disease and even climate. There
activity—through the destruction enough to be catastrophic for an is now a growing commitment from
of natural habitats by deforestation, ecosystem, damaging it beyond many governments across the
urbanization, pollution, and the the point of recovery and so world to use these benefits both
cumulative effects of anthropogenic causing a permanent change responsibly and sustainably. ■
(human-induced) climate change. in its identity, or even its demise.
Humans can also be responsible The fear is that much of the
The Eden Project, in Cornwall, UK,
for the introduction of invasive disturbance caused by human simulates a rain forest ecosystem in
species. Without these external activity has the potential to one of its giant dome greenhouses. The
factors, an ecosystem would cause such permanent damage, domes’ panels are slanted to absorb
maintain its state of equilibrium, particularly when it involves the plenty of light and thermal energy.
and retain a stable identity.

Resistance and resilience

Ecosystems are often strong
enough to withstand some natural
external disturbances and retain
their equilibrium. Some are more
resistant to disturbance than
others, and have adapted to the
particular disturbances normally
associated with their environment.
In some forest ecosystems, for
example, the periodic fires caused
by electrical storms cause only a
minor imbalance in the ecosystem.
Even when severely disrupted
by external disturbances, some
ecosystems have a resilience
that enables them to recover.
However, other ecosystems are



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.
in the prestigious journal Ecology. Lindeman had painstakingly
Raymond Lindeman
Titled “The Trophic-dynamic sampled everything in the lake,
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.

See also: Ecological niches 50–51 ■ Nonconsumptive effects of predators
on their prey 76–77 ■ The food chain 132–133 ■ The ecosystem 134–137
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.


Nelson Hairston (1917–2008)

1949 Aldo Leopold publishes
A Sand County Almanac,
drawing attention to the
ecological impact of hunting
wolves on mountain plant life.


1961 Lawrence Slobodkin,
an American marine
ecologist, publishes The
Growth and Regulation of
Animal Populations, a key
ecology textbook.
1980 Robert Paine describes
the “trophic cascade effect,”
when predators are removed
from an intertidal ecosystem.
1995 The reintroduction of
gray wolves to Yellowstone
National Park sets in motion
a series of ecosystem changes.

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

See also: Predator–prey equations 44–49 ■ The food chain 132–133 ■ The ecosystem 134–137 ■ Energy flow through
ecosystems 138–139 ■ Evolutionarily stable state 154–155 ■ Biodiversity and ecosystem function 156–157

different trophic levels. They he described changes in food

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

Controlling factors
In 1960, the American ecologist Overgrazing brings
Increased rainfall
Nelson Hairston and his colleagues habitat change and loss
encourages vegetation
Frederick Smith and Lawrence of species richness
Slobodkin published a key paper
entitled “Community Structure,
Population Control, and Competition,”
which examined the factors that Herbivore prey eat plants
control populations of animals on


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

Californian yellow bush lupines

are fast growing and invasive. The
plant can upset the ecosystem by
causing elevated nitrogen levels that
attract nonnative species.

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,
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



Robert H. MacArthur
(1930–72), Edward O. Wilson
1948 Canadian lepidopterist
Eugene Munroe suggests a
correlation between island
size and butterfly diversity in
the Caribbean.
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.

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

See also: Evolution by natural selection 24–31 ■ Predator-prey equations 44–49
■ Field experiments 54–55 ■ The ecosystem 134–137

Random dispersal of organisms to islands


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, ❯❯


island of Cuba, because economic
sanctions since the 1960s have
The water acts as meant that fewer boats (and their
Islands are a physical barrier, lizard stowaways) dock in Cuba.
surrounded by water. preventing many species
from entering or leaving. “Island” habitats
In the early 1970s, American
biologist James H. Brown applied
the Wilson–MacArthur model to
“islands” of coniferous forest on 19
mountain ridges in the Great Basin
of California and Utah. The ridges
As species arrive are separated from each other by
Islands are and depart, island a vast sagebrush desert. Brown
ecological systems. populations evolve. discovered that the diversity and
distribution of small mammals
(excluding bats) in the isolated
forests could not be explained in
terms of an equilibrium between
they counted returning species floating vegetation). Most of these colonization and extinction. Some
to observe their recolonization. factors apply to any similar, isolated species had become extinct, but
The Florida Keys experiment habitats, not just actual islands. no new species had arrived for
showed that distance did indeed The impact of humans—who millions of years, so Brown dubbed
play an important role: the farther probably began visiting isolated the mammals “relicts.” A few years
an island was from the mainland, islands in the Pacific at least 3,000 later, his analysis of resident bird
the fewer invertebrates returned to years ago—has sometimes been populations on the ridges revealed
recolonize the area. dramatic. In recent centuries, that new bird species had arrived
New waves of immigration can, people took dogs, cats, goats, from larger, similar forests in the
however, save even faraway island and pigs with them when they Rocky Mountains to the east and
species from extinction. This is colonized islands in the Pacific in the Sierra Nevada to the west.
more likely to happen with certain and elsewhere; inadvertently, they Brown concluded that certain
bird species—which can travel long also carried rats on their boats. species groups—especially those
distances quickly—than with, for On many islands, rats ate the eggs that fly—are more likely to be
example, small mammals. There is of breeding seabirds and the successful immigrants than others.
also the so-called target effect, seeds of endemic plants, some of
where some islands are more which grew nowhere else. On the
favored destinations because of the Galapagos Islands, dogs ate
habitat they provide. Given the tortoise eggs, native iguanas, and
choice of a treeless island and one even penguins. Goats competed
with woodland, a tree-nesting bird with Galapagos tortoises for food
will naturally opt for trees. and wiped out up to five species of Destroying rainforest
plant on the Santiago Islands. for economic gain
Human impact The arrival of humans, however, is like burning a
The key factors influencing the has not always reduced species Renaissance painting
species mix on an oceanic island richness on islands. Researchers to cook a meal.
are its degree of isolation, how long discovered the important role of Edward O. Wilson
it has been isolated, its size, the ship-assisted colonization of
suitability of its habitat, its location islands in the Caribbean. Despite
relative to ocean currents, and its relatively small size, Trinidad,
chance arrivals (for example, for instance, has more species of
organisms washed up on mats of anole lizards than the much larger

The rebirth of
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



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
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

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. ■



iodiversity is shaped
IN CONTEXT globally by new species
appearing and others
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



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
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




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
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,
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

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



Michel Loreau (1954–)
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.

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

See also: Mutualisms 56–59 ■ Keystone species 60–65 ■ The ecosystem
134–137 ■ Organisms and their environment 166 ■ Invasive species 270–273

When researching such issues,

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




Frederic Clements coins

Alexander von Humboldt The earliest forms Andreas Schimper shows the term “climax
sets out for Latin America, of ecological survey how the link between a community” for the
on an expedition that lays are conducted in plant’s physiology and process of stabilization
the groundwork for modern studies of wild fish external conditions over time in ecological
plant geography. by Stephen A. Forbes. is key to plant ecology. communities.

1799 1880S 1898 1916

1845 1895 1899

Pierre-François In showing how plants The changing vegetation

Verhulst formulates an are related to their of sand dunes inspires
equation to predict environments, Johannes Henry Chandler
population growth. Warming unites the fields of Cowles’ idea of
botany and ecology. primary succession.

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.


Robert May applies James H. Brown and Mathew Leibold’s

The concept of the chaos theory to Robert Maurer devise the “metacommunities”
“ecological guild” is predicting rates of concept of macroecology, concept looks at how
introduced in Richard growth and decline in in which ecological populations of a single
B. Root’s thesis on the animal population patterns are analysed species disperse
blue-grey gnatcatcher. dynamics. across large areas. and interact.

1967 1976 1989 2004

1957 1975 1988 1991

The first satellite goes Citizen science John Odling-Smee Ilkka Hanski outlines
into space, heralding enables Fred and Norah suggests that “niche his metapopulation
new technologies in Urquhart to discover constructors” actively theory for species in
wildlife tracking. where monarch change their environment. fragmented habitats.
butterflies go in winter.

Clements argued that these climax that “exploit the same class examples, from ancient oxygen-
communities were bound together of environmental resources,” producing cyanobacteria that altered
and could be thought of as single, regardless of how they do it. the composition of the atmosphere
complex organisms. in prehistoric times, to beavers
Clements was soon challenged New ideas creating wetlands.
by American botanist Henry Many new ideas enriched the
Gleason, who agreed that plant study of ecology in the late 20th Modern methods
communities could be mapped, but and early 21st centuries. The Traditionally, the task of monitoring
argued that since individual plant metapopulation concept was environmental change has been
species have no common purpose, advanced by the Finn Ilkka Hanski, the responsibility of academics
the idea of integrated communities who argued that a population of and professional ecologists, but
was invalid. His view found support a species is made up of differing, millions of interested amateurs now
in the 1950s, in the field studies of dynamic components. One part of provide enormous amounts of raw
Robert Whittaker and the numerical a population may become extinct, data on everything from flowering
research of John Curtis. while another thrives. The thriving dates to butterfly numbers, and
In 1967, American ecologist element may subsequently help from the state of coral reefs to the
Richard Root proposed the idea of reestablish the population that has breeding populations of birds. With
the “guild,” a group of organisms— died out. computer power to quickly process
closely related or otherwise—that In the process, British ecologist vast amounts of data, and with
exploit the same resources. Later, John Odling-Smee argued, so-called Earth’s ecology changing faster
ecologists James MacMahon and “niche-constructor” species create than ever, this “citizen science”
Charles Hawkins refined the a more favorable environment for looks set to become an invaluable
definition of a guild to species themselves—as seen in countless resource for ecology. ■




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.

1750 Carl Linnaeus explains
that the distribution of plants
is determined by climate.
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

Sclater produces a description he distribution, or range, of species’ distributions, but the first
of the zoogeography (the biological communities and to make detailed studies of this
geographical distribution of species varies according to aspect of zoology was the Prussian
animals) of the world’s birds. many factors—including latitude, polymath Alexander von Humboldt,
climate, elevation, habitat, isolation, who traveled to Latin America with
1876 Alfred Russel Wallace
and the species’ characteristics. French botanist Aime Bonpland in
publishes his two-volume book The study of species distribution is 1799. Their five-year expedition
The Geographical Distribution called biogeography. Biogeography laid the basis of plant geography.
of Animals, which becomes is also concerned with how and Humboldt believed observation in
the definitive biogeography why the patterns of distribution situ to be paramount, and used
text for the next 80 years. change over time. sophisticated instruments to make
Early zoologists and botanists meticulous records of both plant
such as Carl Linnaeus were well and animal species, noting all the
aware of geographical variations in factors that could influence the


See also: Modern view of diversity 90–91 ■ Animal ecology 106–113 ■ Island
biogeography 144–149 ■ Big ecology 153 ■ Climate and vegetation 168–169

and New Zealand). The dividing line

between these last two regions,
which runs through Indonesia, is
still known as “Wallace’s Line.”

The unity of nature Plate tectonics

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




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


See also: Distribution of species over space and time 162–163 ■ Metapopulations
186–187 ■ Metacommunities 190–193 ■ Overpopulation 250–251

laid fewer eggs and, as their

fertility rate declined, the rate of
population growth slowly stabilized.

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

Key works

1798 An Essay on the Principle

of Population
1820 Principles of Political
1827 Definitions in Political




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
world—dates back to ancient an environment. By shedding light
Stephen A. Forbes
Greece. Aristotle made copious on the interrelated effects of its
observations of wildlife, and his plant and animal life, they could
BEFORE work laid the foundations for later also help explain the distribution of
1799–1804 Alexander von naturalists. It was not until the species and variations over time. ■
Humboldt pioneers the field 19th century, however, that the
of biogeography in his travels potential of such surveys was
in Latin America. really understood.

1866 German naturalist The new study of ecology

Ernst Haeckel coins the term As naturalists undertook longer
“ecology” to describe the study field trips, the global distribution
of organisms in relation to of species became more apparent,
their environments. and the concept of ecology as a
science gained traction.
1876 After traveling One of the first scientists to
extensively, British naturalist employ ecological methods was
Alfred Russel Wallace American biologist Stephen A.
publishes The Geographical Forbes. In the 1880s, while
Distribution of Animals. studying fish in a Wisconsin lake,
he realized that survey data could
be interpreted to give a picture of
1890s Frederic Clements interactions between different
proposes the notion of species—not just their abundance.
Satellite images enable ecologists to
ecological communities. observe large-scale changes easily. The
Forbes extended the scope of the green areas in this image of the Caspian
1895 In Ecology of Plants conventional survey, combining Sea are evidence of algal growth—the
Johannes Warming describes practical fieldwork with theoretical product of nutrient enrichment.
the impact of the environment
on the distribution of plants. See also: Classification of living things 82–83 ■ Animal ecology 106–113
■ Biodiversity and ecosystem function 156–157



lant ecology examines how
IN CONTEXT plants interact with one
another and with their
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



hat different plants grow
IN CONTEXT in different climates was
likely common knowledge
for as long as agriculture has
Andreas Schimper
existed; many cultures have
traded plants for thousands of years.
BEFORE However, the clear link between a
1737 Carl Linnaeus’s Flora region’s dominant vegetation type
Lapponica includes details of and climate was not categorically
the geographical distribution spelled out until German botanist
of Lapland plants. Andreas Schimper published his
ideas on plant geography in 1898.
1807 Alexander von Humboldt Botanists such as Carl Linnaeus
publishes his seminal Essay and Alexander von Humboldt had
on the Geography of Plants. written about plant distributions in
the 18th and early 19th centuries. “Flowering stones” (Lithops) are
AFTER The widely traveled Humboldt native to southern Africa, their thick,
1916 In Plant Succession: understood that climate was one fleshy leaves well suited to dry, rocky
an Analysis of the conditions. Related species also occur
of the key factors governing where in similar arid habitats in the US.
Development of Vegetation, plants did and did not grow.
Frederic Clements describes Schimper went one step further
how communities of species than Humboldt by explaining plant physiology (the functioning of
are indicators of the climate that similar vegetation types arise plants), it became the foundation of
in which they have matured. under similar climatic conditions the study of plant ecology. Schimper
in different parts of the world. explained that the connection
1968 “The Role of Climate
He then produced a global between the structures of plants
in the Distribution of classification of vegetation zones and the external conditions they
Vegetation,” by American that reflected this observation. faced in different places was the
geographers John Mather Schimper’s 1898 book Plant- key to what he described as
and Gary Yoshioka, explains geography upon a Physiological “ecological plant-geography.”
how temperature and rainfall Basis ran to 870 pages and is one Vegetation was divided into broad
alone are not enough to define of the largest ecology monographs tropical, temperate, arctic,
plant distributions. written by a single author. A mountain, and aquatic zones, then
synthesis of plant geography and subdivided further, according to


See also: Evolution by natural selection 24–31 ■ Ecophysiology 72–73 ■ The ecosystem 134–137 ■ The foundations of plant
ecology 167 ■ Biogeography 200–201 ■ Biomes 206–209

contrivances for regulating the were linked by warm temperatures

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

Adaptations for extremes

Schimper made a close study of
plant physiology—the structures of
plants and how they had adapted to
varying temperature and moisture
conditions. He was particularly
interested in plants growing in
extreme climatic conditions. Salty
environments, for example, require
plants to survive high levels of
soil and water salinity. Schimper
found that vegetation growing on
the coastal mangroves of Brazil, on
Caribbean and Sri Lankan beaches,
and in sulfur-emitting volcanic
craters in Java, were similarly
tolerant to salt.
Schimper also studied how
plants coped in the challenging
conditions of arid environments.
He found that plants growing in
hot, dry places had evolved “varied



Henry Chandler Cowles
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.
1916 Frederic Clements

suggests that communities he Indiana Dunes comprise 15,000 years ago, there would only
settle into a climax, or stable a windswept section of have been bare sand around Lake
equilibrium, at the end of a shifting sand along the Michigan’s shore. Vegetation developed
succession period. in a physical gradient, with sand nearest
southern shore of Lake Michigan, the water and forests farthest back.
US. In 1896, American botanist
1977 Ecologists Joe Connell
Henry Chandler Cowles saw these
and Ralph Slatyer argue that
dunes for the first time, and so decomposing matter created
succession occurs in diverse began his career in the emerging favorable conditions for other plants.
ways, highlighting facilitation field of ecology. Dunes are among As these new plants died, even more
(preparing the way for later some of the planet’s least stable plants could grow.
species), tolerance (of lower landforms, and therefore changes Based on his observations,
resources), and inhibition to their ecology happen relatively Cowles developed the idea of
(resisting competitors). quickly. As Cowles walked among ecological succession, although
the dunes, he noticed that when groundwork for the concept had
certain plants died off, their been laid by earlier naturalists. In


See also: Field experiments 54–55 ■ The ecosystem 134–137 ■ Climax community 172–173 ■ Open community theory
174–175 ■ Biomes 206–209 ■ Romanticism, conservation, and ecology 298

Primary succession
The process of primary succession begins in barren
environments such as bare rock. Hardy species, usually lichens,
appear first and then give way to a stable climax community of
more complex and diverse life forms over hundreds of years.


Small annual shrubs,
plants and Grasses and and shade- Shade-
Bare rock Lichens lichens perennials intolerant trees tolerant trees

Hundreds of years

Pioneer species Intermediate Climax

species community

an 1860 address to members of the include pioneer plants (often lichens resulting from the pioneer species,
Middlesex Agricultural Society, and mosses), followed by grassy shrubs and oak, pine, and hickory
Massachusetts, Henry David plants, small shrubs, and trees. trees will begin to grow. As the
Thoreau had stated: “Though I do trees grow higher, shading out more
not believe that a plant will spring Life after disturbance of the underbrush, the grasses are
up where no seed has been, I have Secondary succession occurs after a replaced by plants able to survive
great faith in a seed.” disturbance that destroys plant life, with low sunlight, and, after around
such as a flood or a fire. The plant 150 years, the forest once more
Growth of an ecoystem life reestablishes itself and develops resembles the prefire community. ■
French geographer Adolphe Dureau into an ecosystem similar to the one
de la Malle is regarded as the first that existed before the disturbance.
person to use the term “succession” The stages of secondary succession
with reference to ecology when he are similar to those of primary
witnessed the progression of plant succession, although the ecosystem
communities after all the trees may start at different points in the I … found indisputable
were removed from a forest. Cowles process, depending on the level of evidence (a) that forests
provided a more formal articulation damage caused by the trigger. succeeded prairie, and
of his ecological succession theory, A common example of secondary (b) that prairie had
in The Ecological Relations of the succession occurs after a wildfire in
succeeded forest.
Vegetation on the Sand Dunes of oak and hickory forests. Nutrients
Lake Michigan, published in 1899. from burned plants and animals
Henry Allan Gleason
American ecologist
In this seminal paper, he proposed provide the right conditions for
the idea of primary succession— growth of annual plants. Pioneer
the gradual growth of an ecosystem grasses soon follow. After several
originally largely devoid of plant life. years, due at least in part to the
The stages of primary succession environmental and soil changes



he term “climax community”
IN CONTEXT In every region, plants was first proposed in 1916
grow and develop by American botanist
through a series Frederic Clements. He used it to
Frederic Clements
of successions. describe an enduring ecological
community that has reached a
BEFORE steady state, such as a naturally
1872 German botanist stable forest of old-growth trees
August Grisebach classifies that has not undergone or been
the world’s vegetation patterns subjected to any unnatural
in relation to climate. At each stage, they changes, such as logging.
become bigger,
1874 British philosopher more complex, and Regional communities
Herbert Spencer suggests that interconnected. In the 19th century, German
the human population can be botanists August Grisebach and
thought of as a giant organism. Oscar Drude were among those
who recognized that patterns of
1899 In the US, Henry Cowles vegetation around the world reflect
proposes that vegetation factors such as climate variations.
develops in stages, a process It was clear, for example, that the
called succession. Eventually the vegetation
typical vegetation in a wet, tropical
takes on the most
AFTER complicated interconnected climate was very different to that in
form the climate will allow. a dry, temperate climate. Then in a
1926 US ecologist Henry
landmark paper in 1899, American
Gleason argues that a climax
botanist Henry Cowles described
community is a coincidental
how plants colonized sand dunes
collection of individuals. around Lake Michigan in stages—
1939 British botanist Arthur or “successions”—of increasing
Tansley suggests there is not size and complexity.
Once a community In an influential book, Plant
a single climax community but reaches this
“polyclimaxes” responding to Succession (1916), Frederic
“climax,” vegetation Clements developed Cowles’s
various factors. stops changing. idea, which he combined with the
biogeographic thinking of the two


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

The Sonora Desert is often seen as

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

German botanists to produce

a theory of the development of
natural communities.
Clements suggested that the
way to understand patterns of
vegetation across the world is
to think in terms of “formations.”
A formation is a large, natural
community of plants dominated
by a range of life forms that reflects
the regional climate. In each region,
plants go through stages or
successions until they reach the
most complex, highly developed
form of vegetation possible. Once
it finally reaches this climax, the
community stabilizes, in what
was later termed a “steady state,” stages of growth, he argued that for identifying such a community,
and stops changing. it can be considered as a single offered by American botanist Frank
Clements then proposed that complex organism. A community Egler in the 1950s, was never
climax communities are bound grows toward a climax in the claimed. Despite the difficulties,
together. Although an ecological same way that an individual ecologists continued to use the
community is made up of a develops through life stages. theory of a climax community to
multitude of plants at different Clements expanded the idea to decide how to respond to invasive
embrace all organisms in a “biome” species that threatened to disturb
that comprised “all the species of an established native community,
plants and animals at home in a and in recent decades Clements’s
particular habitat.” From this, ideas have regained support.
the idea of the ecosystem as a Succession remains a core
“superorganism” later developed. principle of ecology. In general,
For Clements, climates are early succession phases consist of
like genomes, and vegetation A fluctuating process fast-growing and well-dispersed
is like an organism whose Clements’s ideas were challenged species that are replaced by more
characteristics its from the start, although the idea of competitive species. Initially,
genome determines. a “steady state” proved influential ecologists thought that ecological
Christopher Elliott and dominated thinking about succession ended in what they
Philosopher of science ecosystems up until the 1960s. described as the climax phase,
However, scientists realized that when the ecosystem reached a
communities change constantly in stable equilibrium. However, it
response to conditions, and it is is now accepted that ecological
almost impossible to observe a true succession is a dynamic process
climax community. A $10,000 prize that is constantly in flux. ■



IN CONTEXT There is no evidence of

Plants grow according to
KEY FIGURE integrated development
their individual needs
between plants
Henry Allan Gleason
1793 Alexander von Humboldt
uses the word “association” to An ecological They grow randomly,
sum up the range of plant community is not influenced only by the
types in a particular habitat. an organism environmental conditions
1899 In the US, Henry Cowles
states that vegetation develops

in stages, in a process he calls hen American plant could see none of the integration
plant succession. ecologist Frederic proposed by Clements. Instead,
Clements proposed the Gleason believed that groups of
1916 Frederic Clements posits idea of climax communities in 1916, plants were random growths
the idea of a climax community he envisioned the community as a of individuals and species,
as a single organism. superorganism in which all plants responding to local conditions.
AFTER and animals interact to develop the
1935 Arthur Tansley coins community. A year later, American Individual needs
plant ecologist Henry Gleason Gleason maintained that the
the term “ecosystem.”
dismissed the idea; he argued that changes that occur during plant
1947 Robert H. Whittaker plant species have no common succession, as the composition
begins field studies that will purpose but merely pursue their of a community evolves, are
refute Clements’s holistic idea own individual needs. Gleason’s not integrated stages, as in the
of plant communities. hypothesis became known as the development of a single organism.
“open community” theory. The Rather, they are a combination of
1959 John Curtis boosts dispute initiated a debate that still responses from individual species
Henry Gleason’s reputation rages in ecological circles today. as they seek to meet their own
with numerical studies of Gleason did not deny that plant needs within a locality. “Every
prairie plant communities. communities could be mapped and species of plant,” Gleason argued,
their interactions identified, but he “is a law unto itself.” Gleason also


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

Diseases such as American chestnut

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

during the 1930s as holism became

progressively supported by the idea
of the interactive “ecosystem.”
Nonetheless, as ecologists
continued to study the world, they
found more and more flaws in
Clements’s theory. In the 1950s, the
work of American plant ecologists
Robert H. Whittaker and John Curtis
showed how impossible it was to
identify communities as neat units
of holistic theory, and that the real
world was more nuanced and
complex. When it comes to studying
ecosystems in the field, Gleason’s
ideas seem to provide a better fit.
denied that there is any endpoint or need not worry too much about In the ensuing decades, while
climax community; he believed that disturbing the balance of the environmentalists continue to
communities are always changing. natural environment—because champion holistic ideas, ecologists
there is no balance. Gleason’s ideas have also increasingly incorporated
Changing opinions were therefore forgotten in the Gleason’s concepts into their work.
Gleason’s argument with Clements enthusiasm for developing ecology He is now considered to be one
caused quite a stir at the time. as a science. He became so of the most significant figures in
Clements seemed to be creating an frustrated that he gave up ecology 20th-century ecology. ■
overview in which natural patterns
of vegetation were determined by Henry Allan Gleason
clear rules, just as in Newtonian
science the movement of the planets Born in 1882, Henry Gleason classification. With botanist
is dictated by incontrovertible laws. studied biology at the University Arthur Cronquist, he co-wrote a
Clements and his supporters were of Illinois. He held faculty posts definitive guide to the plants of
able to look at the bigger picture, and conducted acclaimed early the northeastern US. He retired
while Gleason was viewed as a ecological research in Sand in 1950 but continued to write
reductionist, myopically intent on Ridge State Forest, Illinois. In and study. He died in 1975.
the details and challenging the the 1920s, Gleason’s theory of
entire idea of ecology as a science individualistic—rather than
controlled by laws. holistic—plant communities was Key works
not accepted by ecologists. This
Gleason appeared to be saying
rejection led Gleason to abandon 1922 “On the Relation between
that there are no patterns in nature:
ecology in the 1930s. He had Species and Area”
it is all random. Worse still, he was long held posts at the New York 1926 “The Individualistic
accused by some of justifying Botanical Garden and became Concept of the Plant
exploitative farming, since his famed for his work on plant Association”
ideas seemed to imply that man


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
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.
1989 In the US, James
MacMahon suggests that it
does not matter how ecological
guild members use resources.
2001 Argentinian ecologists
Sandra Diaz and Marcelo
Cabido propose grouping
species that have a similar
effect on their environment.


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

also consume oak-leaf insects

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



North American migration pathways
Migratory birds
Fred Urquhart (1911–2002), in North America
Norah Urquhart (1918–2009) use paths that can
be divided into four
BEFORE north–south zones,
1883 The Bird Migration called flyways—Pacific,
and Distribution recording Central, Mississippi,
program starts in the US. and Atlantic. Citizen
scientists can play a
1966 The North American key role in recording
Breeding Birds Survey, the birds as they stop
KEY to feed or rest along
conducted by volunteers, the way, during their
begins in Maryland. Pacific flyway flights north in spring
and south in fall.
AFTER Central flyway
2007 The Global Biodiversity
Information Facility (GBIF) Mississippi flyway
launches a global online portal Atlantic flyway
for collecting data on plants
and animals from citizen

scientists and professionals. itizen science is research migration of birds, the Scottish
and observation carried enthusiasts using lighthouses
2010 The eBird online project,
out by nonprofessional around the coast as observation
created in the US in 2002 by individuals, teams, or networks posts. Then, in the early 1880s, the
the Cornell Laboratory of of volunteers, often in partnership idea of collective observation was
Ornithology for volunteers to with professional scientists. It is extended onto a national scale by
report real-time bird sightings, based on an appreciation that the American ornithologist Wells
becomes a global survey. scientific community should be Cooke, who began a project to show
responsive to the environmental arrival dates for migratory North
concerns of society as a whole, and American birds and provide
an understanding that citizens can evidence for migration pathways.
produce reliable scientific evidence Cooke’s project ran until World War
that leads to greater scientific II, gathering 6 million data cards
knowledge. The involvement of on more than 800 bird species and
ordinary people allows research utilizing 3,000 volunteers at its
bodies to accomplish projects that peak. In 2009, the North American
would be far too expensive or time- Bird Phenology Program began
consuming to run otherwise. to digitize the data from the cards,
Butterflies—millions which has provided valuable
upon millions … carpeted Early enthusiasts evidence of changed bird migration
the ground in their While the term “citizen science” dates and routes resulting from
flaming myriads on this is relatively new, dating from the global climate change.
Mexican mountainside. 1980s, the concept and practice The world’s longest-running
Fred Urqhuart of using the public to observe the citizen science survey is the
natural world and record data has Christmas Bird Count (CBC), held
a long pedigree. In the 1870s, each year in the US. Christmas
small groups of ornithologists “side hunts” of birds were a popular
in Germany and Scotland began pastime in many rural districts
to collect reports on the fall of the US in the 19th century,


See also: A system for identifying all nature’s organisms 86–87
■ Big ecology 153 ■ The distribution of species over space and time 162–163

regardless of whether the birds a tagging scheme in an attempt

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

Key works

1960 The Monarch Butterfly

1987 The Monarch Butterfly:
International Traveler


Science should be
dominated by amateurship
instead of money-biased
technical bureaucrats.
Erwin Chargaff
Austro-Hungarian biochemist

their own homes, but simply

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


University of Georgia, uses an Young volunteers at Siyeh Pass,
app, the Marine Debris Tracker, in the state of Montana, record their
to record sightings of debris in sightings of mountain goats for the
high country citizen science project
the ocean. Understanding patterns in the Glacier National Park.
of trash buildup in the world’s seas
helps scientists to track how
it is transported by currents and introduce bias into recordings,
where to concentrate removal efforts such as by the omission of a
for maximum effect. species that cannot be identified.
The advent of new technology Most simple citizen science
has led to a proliferation of citizen tasks, though, require no training,
science projects. Online recording and some other, more complex,
systems allow people to log procedures can be tackled after
sightings of anything from stag basic tuition. People are often
beetles to wildflowers or migrating attracted to citizen science precisely
birds. In the UK, for example, the because they gain new skills in the
Greenspace Information for Greater process. Increasing pressure on
London (GiGL) website, created by Earth’s natural environments and
the National Biodiversity Network, resources creates an ever greater Painting the
allows people to submit records need for data that records presence,
complete picture
online or by phone, adding to a absence, and change in species,
database used by scientists working their habitats, and the wider Citizen scientists are now
to conserve species and habitats. ecosystems. Projects such as the biggest global providers
Zooniverse, the world’s largest of data on the occurrence of
Limitations and potential citizen science platform, help living organisms. Data is
Some ecology research projects fill this need, accumulating data easier than ever to submit
are beyond the reach of untrained from around 1.7 million volunteers and artificial intelligence (AI)
amateurs because they require worldwide. Such projects will be an algorithms can process data
too high a degree of skill, or invaluable resource for conservation in minutes where once it
technology that is too complex organizations, research institutions, would have taken weeks. For
or expensive. People unfamiliar nongovernment agencies, and example, if a person records
with scientific methods may also governments for years to come. ■ sightings of birds coming to
a garden feeder and sends a
report from a phone to Cornell
University’s eBird website, the
information is compared with
Scientific study often relies on the collection
previous data on factors such
of large quantities of data.
as population numbers and
migration routes. More than
390,000 people have submitted
millions of bird sightings to
eBird from nearly 5 million
Volunteer networks are
The more data, the more able to collect vast
locations around the world.
representative the quantities of data, often
This data is fed into the Global
results are of reality. from widespread areas. Biodiversity Information
Facility (GBIF, coordinated in
Denmark), which collects
information on plants, animals,
fungi, and bacteria. GBIF now
The citizen’s network depends contains more than 1 billion
on volunteers. observations, and the number
is growing daily.



haos theory—the idea that
IN CONTEXT predictions are limited by
time and the nonlinear
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



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,
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



IN CONTEXT A species becomes

A species colonizes
extinct in one
KEY FIGURE an empty habitat patch.
habitat patch.
Ilkka Hanski (1953–2016)
1931 In the US, geneticist
Sewall Wright explores the
influence of genetic factors Extinction and colonization are dynamic processes.
on species populations.
1933 In Australia, ecologist
Alexander Nicholson and
physicist Victor Bailey develop A local extinction does not signal
their model of population the extinction of the species.
dynamics to describe the
host–parasite relationship.

1954 In The Distribution and metapopulation is a other places. The species is like a
Abundance of Animals, combination of separate, family whose members have moved
Australian ecologists Herbert local populations of the to different cities yet are still
Andrewartha and Charles same species. The term was coined related. The combined effect of
Birch challenge the idea that by American ecologist Richard many populations may boost the
species populations are Levins in 1969 to describe how long-term survival of the species.
controlled by density alone. insect pest populations rise and fall
on farm fields. Since then, its use Apart but together
AFTER has expanded to cover any species A crucial aspect of metapopulation
2007 American ecologist broken up into local populations in theory is the level of interaction
James Petranka links fragmented habitats, both on land between the separate local
metapopulation theory to and in the oceans. populations. If the level is high,
the metamorphosis stages A particular species of bird, for it is not considered to be a
of amphibians. instance, may be found in separate metapopulation—all the local
populations in a lowland forest, in groups are part of one big
mountain woodlands, and various population. In a metapopulation


See also: Animal ecology 106–113 ■ Clutch control 114–115
■ Island biogeography 144–149 ■ Metacommunities 190–193

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

Key works

1991 Metapopulation
1999 Metapopulation Ecology
2016 Messages from Islands




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


See also: Ecological niches 50–51 ■ The ecosystem 134–137 Organisms and
their environment 166 ■ The ecological guild 176–177

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



Mathew Leibold (1956–)

1917 Arthur Tansley observes
that two species of Galium

plants grow differently in
different soil patches.

METACOMMUNITIES 1934 Georgy Gause develops

the competitive exclusion
principle stating that two
species competing for the
same key resource cannot
coexist for long.
2001 Stephen Hubbell’s
“neutral theory” argues that
biodiversity arises at random.
2006 Mathew Leibold and
fellow American ecologist
Marcel Holyoak refine and
develop the theory of

ne of the limitations
of traditional community
ecology was that it tended
to look at communities purely
locally and take little account of
what happens at different scales
or across different places. Therefore,
over the last few decades, ecologists
have been developing theories of
“meta” communities; the concept
was summed up in 2004 in a key
paper led by American ecologist
Mathew Leibold.
The idea of metacommunities
is linked to that of metapopulations.
While studies of metapopulations
examine the different patches
where populations of the same
species coexist, in metacommunity


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

Mountain goats in Colorado live

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

theory the different patches consist

of entire communities that include
a number of interacting species.

What is a metacommunity?
Metacommunities are essentially
groups or sets of communities.
The communities making up a
metacommunity are separated in
space, but they are not completely
isolated and independent. They feed, shelter, or breed. Differing of seemingly contradictory
interact as various species move types of habitat will influence this observations. One ecologist’s study,
between them. For example, a balance between interlinked and for instance, might look at the way
metacommunity might consist of a independent development. The species live and interact together
set of separate forest communities, theory of metacommunities provides in a small local community. This
spread across a region. The various a framework for studying how and narrowly focused study finds that
species within each patch of forest why variations develop and their competition between species for
habitat interact as an independent impact on biodiversity and resources is a crucial factor in the
community. However, certain population fluctuations. workings of the community.
species, including deer or rabbits, Another study might look at the
may migrate or disperse to another Local versus regional picture across a larger community.
community in the metacommunity, A major advantage of looking at This macro-study discovers that
moving to a different patch of forest communities in this spatial way competition plays virtually no
in search of better opportunities to is that it may help resolve a number part. So which result is correct? ❯❯

Wildlife crossings Many different species cross from one habitat to another. The
naturally between separate idea of providing wildlife with
habitat patches. This movement ways through is not new. For
can be seasonal, as in annual example, fishways for fish to
migrations, or prompted by bypass dams go back centuries.
natural disasters, such as fire or Wildlife crossings—from bridges
flood, or may take place over long for bears in Canada to tunnels
timespans. It creates connections for California’s desert tortoises—
that are often essential for the are becoming an increasingly
health and survival of species and common feature of construction
communities, providing renewal work. Thousands of crossings,
or new resources at pivotal among them bridges, viaducts,
moments. Increasingly, however, and underpasses—often planted
manmade barriers, such as with vegetation—have been
clearances for agriculture, road, built to conserve habitats and
railroads, and urban sprawl, are to avoid fatal collisions between
breaking up this natural interflow animals and vehicles.

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


perch dragonfly pond weed

lake 2
lake 1

heron cormorant
rushes heron frog

lake 3

perch mosquito
rushes algae carp cormorant
lake 4 lake 5

duck algae

In this example of a metacommunity, arrows show

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


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

for a period of time following a

storm, patches of fungal fruiting
bodies that live just a few days
or weeks, and even pitcher plants
that, after dew or rain, provide a
short-lived aquatic home for both
bacteria and insects.

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




Louis Agassiz shows that Vladimir Vernadsky’s

an ice sheet once covered Svante Arrhenius is the book The Biosphere explains
Switzerland, and suggests first to argue that carbon how atmospheric gases are
that an ice age occurred dioxide emissions can lead created by biological
in recent geological history. to global warming. processes.

1840 1896 1926

1869 1912 1935

The father of biogeography Alfred Wegener presents his Arthur Tansley coins the term
Alfred Russel Wallace theory that Earth was once “ecosystem” to describe
reports a clear evolutionary a single landmass from an interdependent
division in fauna species which continents community of biological and
on neighboring islands. drifted apart. nonbiological components.

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


In their book Bioecology, Eugene and Howard Odum James Lovelock’s Gaia
Frederic Clements and describe the living planet hypothesis presents
Victor Shelford popularize as a global collection of Earth as a single,
the idea of the biome. interlocking systems. self-regulating system.

1939 1953 1974

1947 1970 1980

Lesley Holdridge’s The first Earth Day is Luis and Walter

biogeographic classification celebrated, and the idea Alvarez suggest that the
maps 38 life zones where that humans should view mass extinction of the
vegetation is determined Earth holistically dinosaurs was caused
by rain and heat. grows in popularity. by a meteor strike.

seas as a key supporting argument areas at or near the surface of the Almost two centuries earlier Hutton
for evolution. Wallace noted, for Earth where organic life can exist. had articulated a similar idea—that
example, the ocean straits that In 1926, the Russian geochemist biological and geological processes
produced a sharp division between Vladimir Vernadsky explained are interlinked and that Earth could
the flora and fauna of Australasia the biosphere’s close interaction be viewed as a superorganism. In
and Southeast Asia. with the planet’s rock (lithosphere), Hutton’s words, “The globe of this
With a better understanding of water (hydrosphere) and air earth is not just a machine but also
Earth’s biogeography, 20th-century (atmosphere). This in turn led an organized body as it has a
ecologists divided the planet into American biologist Eugene Odum regenerative power.”
biomes—broad communities of flora to advocate a holistic approach to
and fauna that interact in different ecology. Odum argued that it was Heading for extinction?
habitats, such as tropical rain forests not possible to understand a single Life has survived on Earth for
or tundra. Botanist Leslie Holdridge organism, or a group of organisms, billions of years, despite the ravages
refined the concept in 1947 with his without studying the ecosystem in of five mass extinctions. However,
life zone classification, in which he which they live. He described this environmentalists now question
mapped zones based on the two view as “the new ecology.” whether it will survive another.
crucial influences on vegetation: In 1974, British scientist James Indeed, some contend that a sixth
temperature and rainfall. Lovelock advanced the Gaia mass extinction has already started,
hypothesis that the interaction as a result of human activity. Yet,
A “whole Earth” approach of living and nonliving elements if Lovelock’s Gaia theory is correct,
The word “biosphere” was coined in the biosphere reveal Earth to be a it seems likely that the planet will
by Austrian geologist Edward complex, self-regulating system that endure—even if humans and many
Suess in 1875 to signify all the perpetuates the conditions for life. other current life forms do not. ■



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
landforms, plants, and animals. of glaciers, and the landforms they
Louis Agassiz (1807–73)
Supporters of catastrophism argued create, informed these ideas. After
BEFORE that a series of destructive shocks, observing parallel striations in rocks
1795 Scottish geologist James such as the Great Flood described in of the Swiss Alps, German–Swiss
Hutton argues that erratic the Bible, had re-formed the surface geologist Jean de Charpentier (or
boulders (rock fragments that of the planet many times, reshaping Johann von Charpentier) postulated
are different from the existing mountains, lakes, and rivers that glaciers in the Alps had once
underlying rock) in the and wiping out many plant and been more extensive and had
Alps were transported by animal species. In contrast, followers caused the scratches as they moved
moving glaciers. of uniformitarianism contended that and their sediment cut into the rock.
Earth’s features were the result of Geologist Jens Esmark drew similar
1818 In Sweden, naturalist continuous and uniform natural conclusions in Norway.
Göran Wahlenburg publishes processes of erosion, sedimentation
his theory that ice once (the depositing of particles carried Glacier movements
covered Scandinavia. by fluid flows), and volcanism. Swiss zoologist Louis Agassiz
Detailed geological studies developed Charpentier’s and
1824 Danish–Norwegian demonstrated that neither camp Esmark’s ideas further. In 1837,
mineralogist Jens Esmark was right. They established that he proposed that vast sheets of
theorizes that glaciers were ice had once covered much of the
once larger and thicker and northern hemisphere, from the
had covered much of Norway North Pole to the Mediterranean and
and the adjacent seafloor. Caspian coastlines. Agassiz also
undertook some detailed studies
AFTER of glacier movement in Switzerland
1938 Serbian mathematician and published his Études sur les
Milutin Milanković publishes
a theory to explain the
recurrence of ice ages based Animals enter Noah’s ark in a
depiction of the Great Flood described
on changes in Earth’s orbit in the Bible. Catastrophists believed that
around the Sun. the Great Flood was one of the formative
shocks that shaped the geology of Earth.


See also: Evolution by natural selection 24–31 ■ Global warming 202–203
■ The Keeling Curve 240–241 ■ Ozone depletion 260–261 ■ Spring creep 274–279
Receding glaciers
and bird migration
When the last glacial period
began to end, around 26,500
years ago, Earth was much
colder than it is today. Much
of North America and northern
Eurasia was covered with ice
sheets. The environment was
so harsh that most birds
tended to live in subtropical
and tropical regions where
there was more food.
As temperatures began
rising, the ice sheets started
to shrink, uncovering a new
landscape. Ice-free ground
and short, wet summers were
ideal for insects, and birds
began to move in, too, to
take advantage of this food
supply. When days got shorter
in fall, some birds stayed on
for the winter, but others
returned south.
glaciers in 1840. The same year, he Glaciers converge on Piz Argient, The distances flown by
visited geologist William Buckland a mountain in the Swiss Alps. Like birds returning to their homes
others in the Alps, these glaciers were
in Scotland to investigate glacial grew longer as the ice sheets
once much more extensive than they
features there, prompting Scottish are now, and they continue to shrink. retreated farth