KH4119_Unit 01 ES_E110-E120 03/16/05 3:32 PM Page 110

Primates Show Change

across Time
A carpenter walks up to a cabinet she is combination of abilities allows us to use tools
building and places a metal handle in a spot even while we move about. While all primates
she has marked. Holding the handle with her have the ability to grasp objects with their

Y
left hand, she turns the screwdriver with her opposable thumbs to some degree, humans have
right hand to attach the handle to the wood. remarkably nimble fingers and hands. The
Somewhere, 5,000 years ago, a hunter steps biological changes that made the refined use of

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quietly through a thin forest, keeping his sight our hands possible took place over a very long
fixed on a small deer a short distance away. As period of time in human evolution.
he walks, the hunter pulls an arrow from his The fossil record of primates supports the

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quiver and sets it to the bow, ready for a shot idea of biological changes at various rates in
that may win his family several good meals primate history. Today, the primate group
and a hide for clothing. includes such mammals as lemurs,

C
Walking upright has been advantageous for chimpanzees, gorillas, orangutans, and
humans because it frees our fingers and hands humans. Millions of years ago, all primates
to grasp and manipulate objects. This shared a single common ancestor. Those

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Common ancestor of hominoids (apes) and
hominids (upright walkers) Gorillas

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Common ancestor of gorillas, chimpanzees, and
hominids (upright walkers)

Common ancestor of chimpanzees and

hominids (upright walkers)

N
Chimpanzees

O
es
amid
A. r

nsis Hominids
fare
A. a including humans

D 12 mya 9 mya 6 mya 3 mya

Orangutans

present

Figure E2.8 Gorillas, chimpanzees, orangutans, and humans share a common

ancestry. This timeline shows a common interpretation of the general relationship
between some present primates, including humans. Notice that humans are most closely
related to chimpanzees, and that neither humans nor chimpanzees evolved from apes.
Rather, modern apes and humans evolved from a common ancestor.

110 Unit 1 ESSAY: Primates Show Change across Time

KH4119_Unit 01 ES_E110-E120 03/16/05 3:32 PM Page 111
primates that are most closely related shared
additional common ancestors (see Figure E2.8).
Because of her pelvic and leg anatomy, scientists
assigned Lucy to a group of primates called
hominids. Hominids walked upright. Early
hominids had brain cases more similar in size to
apes than to humans. Still, hominid skulls
indicate that their brains were larger for their
body size than those of modern apes are. Lucy
belongs to a group of hominids called
Australopithecus. This group had more
humanlike front teeth, which are considerably
smaller than those in apes. In addition, the
position of the opening that the spinal cord
passes through into the skull indicated an erect,
bipedal posture.
Likewise, Lucys pelvis clearly indicates
that she walked fully upright. By contrast, the
shape of a chimpanzees pelvis indicates that it
is a knuckle-walker. Taken together, Lucys
skeletal remains suggest that her species was
somewhere between apelike ancestors and
modern humans. In 1995, a new species of
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hominid was uncovered in the Lake Turkana
region of Kenya, in East Africa. It had many
of the same features found in Lucy. In
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particular, its leg bones indicate that it stood
upright and walked bipedally. It was named
Australopithecus anamensis, and it was
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Darwin Proposes Descent with Modification
D
Charles Darwin, a 22-year-old English
naturalist, was nervous and excited as he
climbed up the gangplank to the scientific
research ship, HMS Beagle, just after
Christmas in 1831 (see Figure E2.9). Just
imagine how much more excited he might
have been if he had known what his voyage
would mean to modern science. Although
young Charless journey would last only
5 years, the ideas born of that journey would
change forever the way biologists understand
the world.
scientifically determined to be approximately
4.1 million years old. This date makes A.
anamensis more than half a million years older
than Lucys species. Combining this find with
molecular evidence suggests that hominids
split from the common ancestor shared with
chimpanzees about 6 million years ago. The
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fossil A. anamensis moved science closer to
that hypothetical common ancestor (see
Figure E2.8).
P
Anthropologists discovered more
hominid fossils in 2000 and 2002. These
fossils were dated between 5.2 and
O
6.2 million years old. Until more of these
fossils are found and further analysis is
completed, scientists will not know if they
C
represent hominids even closer to that
common ancestor. There seems to be little
doubt that we are getting very close to the
T
root of the hominid family tree.
Other hominids that lived on earth more
recently, such as Homo habilis and Homo erectus,
show a more human appearance. They have a
more vertical face, smaller front teeth, a smaller
lower jaw, and a significantly larger brain case.
These specimens show a gradual change in
form that is consistent with the scientific
explanation that hominids evolved and gave rise
to humans.
The central mission of the Beagle was to
chart sections of the coast of South America.
While the ships crew completed that work,
Darwin spent his time on shore, collecting
thousands of specimens and recording detailed
observations of the interesting and exotic
organisms he encountered. Darwin observed
organisms in such diverse environments as the
jungles of Brazil, the harsh plains of Tierra del
Fuego (near Antarctica), the grasslands of
Argentina, and the heights of the Andes
mountains. Before it was over, the Beagles
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Figure E2.9 Charles

Darwin. Born in 1809,
Darwin studied medicine and
theology to please his father,
but his real interest was in
natural history. Eventually, he
left medical school to enroll
at Cambridge University to

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study natural history, the
term then used for biology.
His mentor, the Reverend

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John Henslow, was a
famous botanist who later
recommended Darwin to the
captain of the HMS Beagle.

voyage took Darwin to the coasts of

Australia, New Zealand, Tasmania, and to

CO
Pacific were clearly distinct. That, perhaps,
was not surprising. But what perplexed

T
several islands in the South Pacific and Atlantic Darwin was the fact that organisms that lived
oceans (see Figure E2.10). in temperate (mild) areas of South America
One of the most important questions that were more similar to organisms living in

O
Darwin pondered during his travels related to tropical areas of South America than to
the geographical distribution of the organisms organisms living in temperate regions of
that he observed. Although similar to Europe. Orchids, army ants, marine iguanas,

N
organisms in Europe, the plants and animals penguins . . . each type of organism was well
that lived in South America and the South suited for the environment in which it lived.

O England Asia

D
Europe
North
Pacific America Atlantic Pacific
Ocean Ocean Africa Ocean
Galpagos Islands equator
South
America
Australia
3000 km
3000 mi
Scale at the Equator
New Zealand

Figure E2.10 Voyage of the HMS Beagle. Darwin sailed around the world on the
Beagle. The route included stops in South America and a stay in the Galpagos Islands.
During the trip, Darwin collected evidence that he later used to support the theory
of evolution.

112 Unit 1 ESSAY: Darwin Proposes Descent with Modification

KH4119_Unit 01 ES_E110-E120 03/16/05 3:32 PM Page 113

Yet each seemed to be related to organisms

living in other parts of that huge continent.
How could one account for both the
similarities and the differences among species.
How could one account for their specific
patterns of geographic distribution?
After he returned to England, Darwin

Y
spent nearly 20 years analyzing his
observations and thinking about their
implications before he published On the

P
Origin of Species. Although this landmark
book established Darwin as the author of the
theory of evolution, Darwin did not use the

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word evolution in the first edition. Instead,
he proposed the concept of descent with
modification. This phrase expressed his view

C
that all organisms on earth are related
through descent from some unknown
ancestral type that lived long ago. This idea

T
helped Darwin develop an explanation for
the diversity of organisms that he had
encountered on his travels and also for the Figure E2.11 Alfred Russel Wallace.
patterns in geographical distributions that he

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had observed. The concept of descent with
modification is described in more detail in
the next essay, Evolution by Natural Selection.
Although Wallace independently reached
many of the same conclusions that Darwin
reached, he seldom is acknowledged for his
contributions to the theory of evolution.

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The publication of Darwins book
represents an interesting twist in the history of
science. If Alfred Russel Wallace, a young
scientist working in the East Indies, had not
Darwin wrote in 1844 but never published.
This decision preserved Darwins claim to
these ideas, but Darwin still had to publish his

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written to Darwin, Darwin might not have full results. Motivated now by the concern that
published the book when he did. In his letter other scientists were about to reach the same
to Darwin, Wallace enclosed a draft of a conclusions that he had reached, Darwin

D
scientific paper that described a theory of
evolution. His theory was less detailed than
Darwins theory, but almost identical in basic
outline. Darwin was shocked to learn that
other scientists not only were thinking about
these ideas but also might be able to publish
them before he did. Nevertheless, he behaved
with integrity, acknowledging the excellence
and importance of Wallaces work and
forwarding Wallaces paper to another noted
scientist for public presentation. Knowing of
Darwins nearly 20 years of study, the scientist
decided to present Wallaces paper with an
excerpt from an essay about evolution that
worked feverishly for almost a year, finally
completing On the Origin of Species in 1859.
Although Wallace published his paper
first, Darwins explanation of evolution was
more detailed and contained more extensive
supporting evidence than Wallaces paper.
Darwins journals also demonstrated that he
had developed the central core of his ideas
more than 15 years before reading Wallaces
paper. Consequently, Darwin is known as
the main author of the theory of evolution,
and even Wallace thought that Darwin
deserved the credit. (Wallace is pictured in
Figure E2.11.)

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Evolution by Natural Selection
Even before Darwins work, several individuals
already had proposed the basic idea of
evolution, the concept that differences among
species are the result of changes across time.
Unlike these earlier scientists, however,
Darwin provided not only a logical argument
to support evolution, but also clear evidence
for his key points. Even more important,
Darwin proposed a mechanism, that is, a way
by which a new species could eventually
appear from ancestral forms.
Although he was unable to explain the
source of variation in organisms, Darwin
observed that the individual members of any
species show a great deal of variation in their
characteristics. (Later, scientists found the
source of variation to be genetics.) They may
T
show differences in size, coloration, strength,
behavior, and many other features. Some of
these characteristics are passed on from
O
parents to offspring. For centuries, in fact,
plant and animal breeders have bred
N
cauliflower kale
O
D
cabbage brussels
sprouts
Figure E2.12 Six types of kale. These kale plants differ primarily in the part of the plant that stores the
most starch.
114 Unit 1 ESSAY: Evolution by Natural Selection
organisms to emphasize or increase certain
prized characteristics. Plant breeders, for
example, try to improve disease resistance by
selecting plants with high resistance and
mating them together. They hope to obtain
Y
offspring with even higher resistance.
Likewise, horse breeders may select mating
P
pairs to try to increase speed, stamina, or both.
Sometimes these techniques can result in
significant differences from one breeding
group to another. Figure E2.12 illustrates six
O
different types of vegetables that humans eat.
All six plants are of the same species. Humans
C
have bred them for different agricultural uses
and to suit different human tastes.
If humans can bring about change in a
population of organisms by this type of
artificial selective breeding, Darwin reasoned
that perhaps selection in nature also could
bring about change. Such change eventually
might result in the production of new species.
A species is a group of organisms whose
broccoli
kohlrabi
KH4119_Unit 01 ES_E110-E120 03/16/05 3:32 PM Page 115

List of Mayrs points

Fact 1: All species have such great potential to produce large numbers of offspring
that their population size would increase exponentially if all individuals that are born
would reproduce successfully.

Y
Fact 2: Except for seasonal fluctuations, most populations are normally stable in size.

Fact 3: Natural resources are limited, and in a stable environment, they remain rela-

P
tively constant.

Inference 1: Because more individuals are produced than the available resources can
support, and the population size remains stable, there must be a fierce struggle for

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existence among the individuals of a population. This results in the survival of only a
part, often a very small part, of the offspring of each generation.

C
Fact 4: No two individuals in a population are exactly the same; rather, every popula-
tion displays an enormous variety of characteristics.

Fact 5: Much of this variation can be inherited.

T
Inference 2: Survival in the struggle for existence is not random but depends in part
on the characteristics that the surviving individuals inherited. This unequal survival is
a process of natural selection that favors individuals with characteristics that fit them

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best in their environment.

Inference 3: Over the generations, this process of natural selection will lead to a
continuing gradual change in populations, that is, to evolution and the production of

N
new species.

O
Figure E2.13 The logic of the theory of natural selection. Ernst Mayr has

D
summarized the logic of Darwins theory by extracting the major points of evidence and
inference. (This list is adapted from E. Mayr, The Growth of Biological Thought: Diversity,
Evolution, and Inheritance, Cambridge, MA: Harvard University Press, 1982.)

members very closely resemble each other
because they share many key characteristics.
Despite obvious differences in superficial
appearance, all humans are one species
because we share many fundamental
characteristics. Organisms belonging to
different species usually vary from one
another in numerous key characteristics.
For sexually reproducing organisms, one
characteristic that separates species is the
ability to mate and produce fertile offspring.
A horse and a donkey, for instance, can mate
and produce offspring called mules. But
because horses and donkeys belong to two
different (although very closely related)
species, these offspring mules are sterile.
How would the process of natural
selection take place? One way to trace

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KH4119_Unit 01 ES_E110-E120 03/16/05 3:32 PM Page 116

Darwins thinking is to consider carefully the
information displayed in Figure E2.13. This
summary was proposed by Ernst Mayr, a
noted biologist at Harvard University. He
introduced a way to understand the mix of
evidence and inference that underlies the
theory of evolution. Let us follow the
out among the members of a natural
population:
Individual organisms within a
population are different in particular
characteristics from one another.
Most, although not all, of this variation
can be inherited from one organism to

Y
argument, point by point, and try to
the next.
understand some of the thinking that led
Darwin to his conclusions.

P
The first three facts in Figure E2.13
describe the situation that inevitably exists in
a
any environment:

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In the absence of limits to population
growth, organisms reproduce very
rapidly.

C
Nevertheless, our observations tell us
that most populations in the wild are
relatively constant in size.

T
Our observations also tell us that the
resources available in any natural
environment are limited. b

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One important factor that influenced
Darwins views was an essay written in 1798
by Thomas Malthus, a member of the English

N
clergy. Malthus argued that much of the
human suffering that we see on earth is the
inescapable result of the tendency of the
human population to grow beyond the

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resources available to support it. This capacity
for overpopulation seemed to Darwin to be a
characteristic of every population. It led c

D
Darwin to infer that individual organisms
within a population must face a struggle for
survival, and that only a few individuals need
to survive to pass on their characteristics from
one generation to the next. The rest fail to
develop; die of starvation, predation, or other
causes before they reproduce; or do not
reproduce for other reasons. The effect of this
loss of reproductive individuals from one Figure E2.14 Examples of
adaptations. (a) snake caterpillar
generation to the next is that wild populations
(b) flicker (c) red Irish lord fish Can you
tend to be relatively stable in size. identify the adaptations possessed by these
The remaining two facts suggested to organisms? How might each adaptation
Darwin how this struggle for survival is played benefit the organism?

116 Unit 1 ESSAY: Evolution by Natural Selection

KH4119_Unit 01 ES_E110-E120 03/16/05 3:32 PM Page 117
From these observations, Darwin inferred
that the outcome of the struggle for existence
depends, to a certain extent, on the
characteristics that an organism inherits.
There are individuals whose inherited
characteristics best equip them to survive in a
particular environment. These individuals will
be most likely to reproduce and to leave
behind offspring with the same beneficial
traits. We call these beneficial traits
adaptations (Figure E2.14). By contrast,
those organisms with characteristics that
make it more difficult to survive and
reproduce in an environment would be less
likely to survive and to reproduce. As a result,
their particular characteristics would be less
likely to be passed on to surviving offspring.
For example, suppose a population of
rabbits lived in a moderately warm climate.
Within that population, a small proportion of
the individuals might have thick fur. If this
thick fur resulted from a difference in their
genetic material as compared with that of
O
other rabbits living in the same area, then this
difference might be passed on to their
offspring. Imagine first that the thicker fur
N
is a disadvantage to the rabbits in this
environment because they overheat when
they are chased by coyotes. As a consequence,
these thick-furred rabbits do not survive and
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reproduce as well as the rabbits with thin fur.
You probably can see, in this case, that thick
fur would not be a very common trait in this
D
population.
Now imagine that, through a period of
several decades, the climate in this
environment gets colder. In the colder climate,
the rabbits with the thicker fur now would
have an advantage. That is, the thick fur
would be an adaptation that helps them
survive the periods of cold weather. All other
things being equal, the thick-furred rabbits
would be more likely to survive long enough
to reproduce and pass on their characteristics
than the thin-furred rabbits. As Figure E2.15
shows, after many generations, the number of
rabbits with thick fur might increase in the
Warm Climate Cold Climate
relative number of rabbits in population
(40 years later)
Y
P
thin fur thick fur thin fur thick fur
O
Figure E2.15 Characteristics of a
population change due to natural
C
selection.
population. This is the process of natural
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selection.
Although Darwins understanding of
selective breeding led him to propose his theory
of natural selection, there is a key difference
between selective breeding and natural
selection. In selective breeding, humans choose
how to allow organisms to reproduce in order to
pass on desirable characteristics. In natural
selection, however, each organisms ability to
meet the challenges of survival and
reproduction in a natural setting determines the
mix of characteristics that will be transmitted to
the next generation.
In such a setting, many pressures affect
survival and reproduction. As you saw in the
rabbit example, changes in the environment
can alter the balance of life so that some
characteristics are more beneficial than others.
Sometimes these changes are catastrophic, like
the effect a volcanic eruption might have on
both local and very distant populations of
organisms.
Competition among organisms for
limited resources also acts as a selective
pressure. For example, a plant needs sunlight
to grow and sufficient soil to support its roots
and provide water and minerals. The plant
must cope with its environment to survive and
ESSAY: Evolution by Natural Selection Unit 1 117
KH4119_Unit 01 ES_E110-E120 03/16/05 3:33 PM Page 118
produce seeds for the next generation. The
plant, however, is not alone. Nearby plants
may grow faster than this plant and may block
the sunlight, or they may have longer roots
that drain off limited water. In that way, these
other plants may compete with the first plant
for survival. A difference in the ability of
organisms to compete in an environment with
T
limited resources is one of the selective
pressures that can bring about evolution.
Predation is another pressure that many
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organisms face (see Figure E2.16). Living
long enough to reproduce requires that the
predator be successful in locating and catching
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prey. The prey, however, must escape
predation until it is able to reproduce. In a
predator/prey relationship, it may appear that
the predator has all the advantages. Not so.
O
Think of a lion whose jaw is broken by a well-
aimed kick from a gazelle. The lion will die
slowly of starvation because it can no longer
D
eat. On the other hand, the gazelle that is not
alert enough or strong enough to avoid the
lions jaws also will die, in this case, rather
suddenly and dramatically.
What determines an organisms ability
to survive and reproduce in the face of these
pressures? Generally, it is not one characteristic
or two, but a mix of characteristics that helps
the organism adapt to a particular environment.
Organisms, however, do not acquire these
characteristics to help them survive. Rather,
individuals inherit their set of characteristics
118 Unit 1 ESSAY: Evolution by Natural Selection
Figure E2.16
The predator/prey
relationship
imposes selective
pressure on living
organisms.
P Y
from their parents. Those individuals in a
O
population who have the characteristics that
make them best suited to their environment
are most likely to survive and contribute their
C
characteristics to the next generation.
It is important to realize that individual
organisms do not evolve. Rather, populations
evolve. Darwins theory of evolution by natural
selection proposes that sometimes the
characteristics of a population can change
so dramatically that eventually the population
becomes recognized as a distinct species,
different from the ancestral form. Here we
return to Darwins basic idea: descent by
modification. Descent from a common
ancestor provides a powerful explanation for
the similarities that we see among related
forms of life. Modification through variation
and natural selection provides a powerful
explanation for their differences.
We cannot overestimate the importance
to modern biology of the theory of evolution
by natural selection. Perhaps Darwins greatest
contribution was in giving biologists a way to
understand the enormous diversity that we see
in living systems. The theory of evolution
explains the origin of this diversity and helps
us to understand the relationships that exist
among modern species and between modern
species and their ancestral forms. As most
biologists will tell you, these aspects of biology
do not make much sense except in the context
of evolution.
KH4119_Unit 01 ES_E110-E120 03/16/05 3:33 PM Page 119

Just a Theory?

What do we mean when we say that we

My theory is that this
have a theory about something? Usually man's son, Niles, ran him Do you think Sergeant
we mean that we have an idea about down to gain the

Y
knows that Niles is only
inheritance money. 3 years old?
how to explain something. We may
suggest a theory about who killed the

P
butler in the late-night thriller. Or we
may have a theory about why Ms.
Figueroa always looks so tired when she

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delivers the mail on Monday morning.
Or we may have a theory about what
the coach of the local baseball team said

C
to his pitcher after they lost last nights
big game.
These theories that we develop
about things are often tentative. That

T
is, when we say that we have a theory
about something, we may mean that we
Figure E2.17 The word theory often means
dont have much evidence. If we dont

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different things to different people.
have much evidence, other people dont
usually agree with or share our
theories. Nevertheless, it is fun to pieces such as electrons and protons, the

N
suggest clever ideas. general outline of his theory still stands.
In contrast, scientists use the word theory in Another example is the cell theory, which
a very different way than we do. Scientific is the understanding that complex living
theories are explanations that are extremely well organisms are comprised of small building

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accepted by the scientific community because a blocks called cells. This is an example of a
variety of strong evidence supports them. biological theory that is supported by an
A good example of a scientific theory is enormous body of evidence.

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the atomic theory, an explanation developed in
1803 by the British teacher and chemist John
Dalton. You may already be familiar with one
of the most important ideas of Daltons
atomic theory: All matter (or things) in the
universe is composed of tiny particles called
atoms. This idea was revolutionary at the time way most nonscientists do.
that Dalton first suggested it. But since then,
physicists have accumulated an enormous
body of evidence to support it. Part of that
evidence is the result of our modern ability to
detect, to manipulate, and even to subdivide
atoms. Even though Dalton did not predict
that atoms could be subdivided into smaller
Despite the evidence that has accumulated
in favor of the atomic and cellular theories,
and despite the fact that no modern scientist
questions most of these basic principles, we still
refer to these ideas as theories. Clearly, scientists
use the word theory very differently from the
To understand how an explanation
becomes known as a scientific theory, we have
to consider first the meaning of the term
hypothesis. You may have learned already
that a hypothesis is a trial idea about
something. During research, scientists
develop hypotheses to explain the reasons

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underlying an event or a phenomenon. A
hypothesis can also explain the relationships
that scientists think they see among objects,
events, or processes. As you will see in Unit 2,
predictions made from such hypotheses are
useful because they can be tested. Testing our
hypotheses allows us to distinguish those trial
ideas that adequately explain what we study
from those that do not.
You probably can see, then, that the
explanations we develop for things and events
around us actually are hypotheses, not scientific
theories. As hypotheses, these trial ideas or
tentative explanations can be tested. In scientific
terms, the police who investigate the death of
the butler do so by systematically testing a
whole set of hypotheses, not theories.
By contrast, a scientific theory is a
hypothesis that already has been extensively
T
tested and is supported by a large body of
observations and evidence. A good theory
explains data that we already know and relates
and explains additional data as they become
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known. In fact, a good theory also predicts new
data and suggests new relationships that we
may not already have recognized.
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Why do scientists call Darwins
explanation of descent with modification the
theory of evolution instead of the hypothesis of
evolution? First, it is considered a theory
O
because of the enormous amount of evidence
that suggests it is a correct explanation. As you
already have seen, this evidence spans a wide
D
range of different scientific fields, from
anatomy to geology to embryology to physical
anthropology. Second, it is considered a
theory because it explains both the evidence
that Darwin saw and recorded, as well as the
new data that we continue to collect. It also
successfully predicts new phenomena. That is,
Darwins explanation for biological change
across time continues to be supported, even by
evidence collected more than 100 years after
his work. For those reasons and others, this
explanation is almost universally accepted by
scientists around the world.
120 Unit 1 ESSAY: Just a Theory?
Y
Actually, Sergeant, I think you mean
OK, I guess my
your hypothesis is wrong. Wed better
P
theory is wrong.
come up with a new hypothesis about
who killed Mr. Smith.
O
C
Figure E2.18 When used by scientists, the
word theory means an explanation that is
supported by evidence and well accepted by
other scientists.
Does this wide acceptance of Darwins
work mean that Darwin explained everything
there was to explain about evolution or that
every part of his explanation is correct? No, not
necessarily. Darwin was not able to explain
exactly how characteristics passed from one
generation to the next. He did not include in his
theory an explanation of the relationship
between natural selection and genetic
inheritance. However, this omission does not
make Darwins ideas incorrect. Since Darwins
time, biologists have added information on
natural selection and genetic inheritance to the
theory of evolution. In fact, the addition of
information to Darwins basic proposal
illustrates the power of a sound scientific theory.
As new information about inheritance became
available, it supported, rather than contradicted,
the original explanation.
So, when we talk about explanations for
things in the world around us, we need to be
careful. We need to decide whether are we
talking about a theory (as we typically use the
term) or are we talking about a theory (as
scientists use the term). The difference
between the two is quite large, indeed.