22

Descent with
(Figure 22.1). Tilting head-downward, the beetle faces into
the winds that blow fog across the dunes. Droplets of mois-
ture from the fog collect on the beetle’s body and run down
into its mouth.
Interesting in its own right, this headstander beetle is also
a member of an astonishingly diverse group: the more than
350,000 species of beetles. In fact, nearly one of every five

Modification: known species is a beetle. These beetles all share similar fea-
tures, such as three pairs of legs, a hard outer surface, and two

A Darwinian pairs of wings. But they also differ from one another. How did
there come to be so many beetles, and what causes their sim-

View of Life
ilarities and differences?
The headstander beetle and its many close relatives illus-
trate three key observations about life:

• the striking ways in which organisms are suited for life in

their environments*
• the many shared characteristics (unity) of life
• the rich diversity of life

A century and a half ago, Charles Darwin was inspired to de-

velop a scientific explanation for these three broad observa-
tions. When he published his hypothesis in The Origin of
Species, Darwin ushered in a scientific revolution—the era of
evolutionary biology.
For now, we will define evolution as descent with modifica-
tion, a phrase Darwin used in proposing that Earth’s many
species are descendants of ancestral species that were different
from the present-day species. Evolution can also be defined
more narrowly as a change in the genetic composition of a
population from generation to generation, as discussed fur-
ther in Chapter 23.
Whether it is defined broadly or narrowly, we can view evo-
! Figure 22.1 How can this beetle survive lution in two related but different ways: as a pattern and as a
in the desert, and what is it doing?
process. The pattern of evolutionary change is revealed by data
from a range of scientific disciplines, including biology, geol-
EVOLUTION
ogy, physics, and chemistry. These data are facts—they are ob-
KEY CONCEPTS servations about the natural world. The process of evolution
22.1 The Darwinian revolution challenged traditional consists of the mechanisms that produce the observed pattern
views of a young Earth inhabited by unchanging of change. These mechanisms represent natural causes of the
species natural phenomena we observe. Indeed, the power of evolu-
22.2 Descent with modification by natural selection tion as a unifying theory is its ability to explain and connect
explains the adaptations of organisms and the a vast array of observations about the living world.
unity and diversity of life As with all general theories in science, we continue to test
22.3 Evolution is supported by an overwhelming our understanding of evolution by examining whether it can
amount of scientific evidence account for new observations and experimental results. In this
and the following chapters, we’ll examine how ongoing dis-
OVERVIEW
coveries shape what we know about the pattern and process
Endless Forms Most Beautiful of evolution. To set the stage, we’ll first retrace Darwin’s quest
to explain the adaptations, unity, and diversity of what he
In the coastal Namib desert of southwestern Africa, a land called life’s “endless forms most beautiful.”
where fog is common but virtually no rain falls, lives the bee-
tle Onymacris unguicularis. To obtain the water it needs to sur- *Here and throughout this book, the term environment refers to other organ-
vive, this insect relies on a peculiar “headstanding” behavior isms as well as to the physical aspects of an organism’s surroundings.

452 UNIT FOUR Mechanisms of Evolution

 1809
Lamarck publishes his
hypothesis of evolution.

1798
Malthus publishes
”Essay on the Principle
of Population.”

1812 1858
1795 Cuvier publishes his While studying species in the Malay
Hutton proposes extensive studies of 1830 Archipelago, Wallace (shown above
his principle of vertebrate fossils. Lyell publishes in 1848) sends Darwin his hypothesis
gradualism. Principles of Geology. of natural selection.

1790 1870
1809 1859
Charles Darwin 1831–36
On the Origin of Species
is born. Darwin travels is published.
around the world
on HMS Beagle.
1844
Darwin writes his
essay on descent
with modification.

Marine iguana in the Galápagos Islands

! Figure 22.2 The intellectual context of Darwin’s ideas.

22.1
changing). Through his observations of nature, Aristotle rec-
CONCEPT
ognized certain “affinities” among organisms. He concluded
that life-forms could be arranged on a ladder, or scale, of in-
The Darwinian revolution creasing complexity, later called the scala naturae (“scale of na-
challenged traditional views ture”). Each form of life, perfect and permanent, had its
of a young Earth inhabited allotted rung on this ladder.
These ideas were generally consistent with the Old Testa-
by unchanging species ment account of creation, which holds that species were indi-
What impelled Darwin to challenge the prevailing views about vidually designed by God and therefore perfect. In the 1700s,
Earth and its life? Darwin’s revolutionary proposal developed many scientists interpreted the often remarkable match of or-
over time, influenced by the work of others and by his travels ganisms to their environment as evidence that the Creator
(Figure 22.2). As we’ll see, his ideas had deep historical roots. had designed each species for a particular purpose.
One such scientist was Carolus Linnaeus (1707–1778), a
Swedish physician and botanist who sought to classify life’s
Scala Naturae and Classification of Species
diversity, in his words, “for the greater glory of God.” Linnaeus
Long before Darwin was born, several Greek philosophers sug- developed the two-part, or binomial, format for naming
gested that life might have changed gradually over time. But species (such as Homo sapiens for humans) that is still used to-
one philosopher who greatly influenced early Western sci- day. In contrast to the linear hierarchy of the scala naturae,
ence, Aristotle (384–322 BCE), viewed species as fixed (un- Linnaeus adopted a nested classification system, grouping

CHAPTER 22 Descent with Modification: A Darwinian View of Life 453

similar species into increasingly general categories. For exam- idea of evolution. To explain his observations, he advocated
ple, similar species are grouped in the same genus, similar gen- catastrophism, the principle that events in the past oc-
era (plural of genus) are grouped in the same family, and so on curred suddenly and were caused by mechanisms different
(see Figure 1.14). from those operating in the present. Cuvier speculated that
Linnaeus did not ascribe the resemblances among species each boundary between strata represented a catastrophe,
to evolutionary kinship, but rather to the pattern of their cre- such as a flood, that had destroyed many of the species living
ation. A century later, however, Darwin argued that classifica- at that time. He proposed that these periodic catastrophes
tion should be based on evolutionary relationships. He also were usually confined to local regions, which were later re-
noted that scientists using the Linnaean system often grouped populated by different species immigrating from other areas.
organisms in ways that reflected those relationships. In contrast, other scientists suggested that profound
change could take place through the cumulative effect of slow
Ideas About Change over Time but continuous processes. In 1795, Scottish geologist James
Darwin drew from the work of scientists studying fossils, the Hutton (1726–1797) proposed that Earth’s geologic features
remains or traces of organisms from the past. Many fossils are could be explained by gradual mechanisms still operating to-
found in sedimentary rocks formed from the sand and mud day. For example, he suggested that valleys were often formed
that settle to the bottom of seas, lakes, swamps, and other by rivers wearing through rocks and that rocks containing ma-
aquatic habitats (Figure 22.3). New layers of sediment cover rine fossils were formed when sediments that had eroded from
older ones and compress them into superimposed layers of the land were carried by rivers to the sea, where they buried
rock called strata (singular, stratum). The fossils in a particu- dead marine organisms. The leading geologist of Darwin’s
lar stratum provide a glimpse of some of the organisms that time, Charles Lyell (1797–1875), incorporated Hutton’s think-
populated Earth at the time that layer formed. Later, erosion ing into his principle of uniformitarianism, which stated
may carve through upper (younger) strata, revealing deeper that mechanisms of change are constant over time. Lyell pro-
(older) strata that had been buried. posed that the same geologic processes are operating today as
Paleontology, the study of fossils, was developed in in the past, and at the same rate.
large part by French scientist Georges Cuvier (1769–1832). In Hutton and Lyell’s ideas strongly influenced Darwin’s
examining strata near Paris, Cuvier noted that the older the thinking. Darwin agreed that if geologic change results from
stratum, the more dissimilar its fossils were to current life- slow, continuous actions rather than from sudden events,
forms. He also observed that from one layer to the next, then Earth must be much older than the widely accepted age
some new species appeared while others disappeared. He in- of a few thousand years. It would, for example, take a very
ferred that extinctions must have been a common occur- long time for a river to carve a canyon by erosion. He later
rence in the history of life. Yet Cuvier staunchly opposed the reasoned that perhaps similarly slow and subtle processes
could produce substantial biological change. Darwin was not
the first to apply the idea of gradual change to biological
1 Rivers carry sediment into evolution, however.
aquatic habitats such as seas and
swamps. Over time, sedimentary
rock layers (strata) form under
Lamarck’s Hypothesis of Evolution
water. Some strata contain fossils. During the 18th century, several naturalists (including Darwin’s
grandfather, Erasmus Darwin) suggested that life evolves as
environments change. But only one of Charles Darwin’s pred-
ecessors proposed a mechanism for how life changes over
2 As water levels
change and the time: French biologist Jean-Baptiste de Lamarck (1744–1829).
bottom surface is Alas, Lamarck is primarily remembered today not for his
pushed upward, the
strata and their visionary recognition that evolutionary change explains pat-
fossils are exposed. terns in fossils and the match of organisms to their environ-
ments, but for the incorrect mechanism he proposed to
explain how evolution occurs.
Younger stratum
with more recent Lamarck published his hypothesis in 1809, the year Darwin
fossils was born. By comparing living species with fossil forms,
Lamarck had found what appeared to be several lines of de-
Older stratum
with older fossils scent, each a chronological series of older to younger fossils
leading to a living species. He explained his findings using two
! Figure 22.3 Formation of sedimentary strata with principles that were widely accepted at the time. The first was
fossils. use and disuse, the idea that parts of the body that are used

454 UNIT FOUR Mechanisms of Evolution

 CONCEPT
22.2
Descent with modification
by natural selection explains
the adaptations of organisms
and the unity and diversity of life
As the 19th century dawned, it was generally thought that
species had remained unchanged since their creation. A few
clouds of doubt about the permanence of species were begin-
ning to gather, but no one could have forecast the thundering
storm just beyond the horizon. How did Charles Darwin be-
come the lightning rod for a revolutionary view of life?

Darwin’s Research
! Figure 22.4 Acquired traits cannot be inherited. This
Charles Darwin (1809–1882) was born in Shrewsbury, in west-
bonsai tree was “trained” to grow as a dwarf by pruning and shaping.
However, seeds from this tree would produce offspring of normal size. ern England. Even as a boy, he had a consuming interest in na-
ture. When he was not reading nature books, he was fishing,
extensively become larger and stronger, while those that are hunting, and collecting insects. Darwin’s father, a physician,
not used deteriorate. Among many examples, he cited a giraffe could see no future for his son as a naturalist and sent him to
stretching its neck to reach leaves on high branches. The sec- medical school in Edinburgh. But Charles found medicine
ond principle, inheritance of acquired characteristics, stated that boring and surgery before the days of anesthesia horrifying.
an organism could pass these modifications to its offspring. He quit medical school and enrolled at Cambridge University,
Lamarck reasoned that the long, muscular neck of the living intending to become a clergyman. (At that time in England,
giraffe had evolved over many generations as giraffes many scholars of science belonged to the clergy.)
stretched their necks ever higher. At Cambridge, Darwin became the protégé of the Reverend
Lamarck also thought that evolution happens because organ- John Henslow, a botany professor. Soon after Darwin gradu-
isms have an innate drive to become more complex. Darwin ated, Henslow recommended him to Captain Robert FitzRoy,
rejected this idea, but he, too, thought that variation was who was preparing the survey ship HMS Beagle for a long voy-
introduced into the evolutionary process in part through in- age around the world. Darwin would pay his own way and
heritance of acquired characteristics. Today, however, our un- serve as a conversation partner to the young captain. FitzRoy,
derstanding of genetics refutes this mechanism: Experiments who was himself an accomplished scientist, accepted Darwin
show that traits acquired by use during an individual’s life are because he was a skilled naturalist and because they were of
not inherited in the way proposed by Lamarck (Figure 22.4). the same social class and close in age.
Lamarck was vilified in his own time, especially by Cuvier,
who denied that species ever evolve. In retrospect, however,
The Voyage of the Beagle
Lamarck did recognize that the match of organisms to their Darwin embarked from England on the Beagle in December
environments can be explained by gradual evolutionary 1831. The primary mission of the voyage was to chart poorly
change, and he did propose a testable explanation for how known stretches of the South American coastline. While the
this change occurs. ship’s crew surveyed the coast, Darwin spent most of his time
on shore, observing and collecting thousands of South Amer-
CONCEPT CHECK 22.1 ican plants and animals. He noted the characteristics of plants
1. How did Hutton’s and Lyell’s ideas influence Darwin’s and animals that made them well suited to such diverse envi-
thinking about evolution? ronments as the humid jungles of Brazil, the expansive grass-
2. MAKE CONNECTIONS In Concept 1.3 (pp. 19–20), lands of Argentina, and the towering peaks of the Andes.
you read that scientific hypotheses must be testable Darwin observed that the plants and animals in temperate
and falsifiable. Applying these criteria, are Cuvier’s regions of South America more closely resembled species liv-
explanation of the fossil record and Lamarck’s hy- ing in the South American tropics than species living in tem-
pothesis of evolution scientific? Explain your answer perate regions of Europe. Furthermore, the fossils he found,
in each case. though clearly different from living species, were distinctly
South American in their resemblance to the living organisms
For suggested answers, see Appendix A.
of that continent.

CHAPTER 22 Descent with Modification: A Darwinian View of Life 455

Darwin in 1840, HMS Beagle in port
after his return
from the
voyage
Great
Britain EUROPE
NORTH
AMERICA
ATLANTIC
OCEAN
The AFRICA
Galápagos PACIFIC
Pinta OCEAN
Islands
Genovesa Equator
Marchena
Equator SOUTH Malay Archipelago
AMERICA PACIFIC
Santiago
Daphne OCEAN
Chile Brazil

Andes Mtns.
Islands
Pinzón AUSTRALIA
Fernandina PACIFIC
Isabela Santa Santa OCEAN Cape of
Cruz Fe San Argentina Good Hope
Cristobal
0 20 40 Tasmania
Florenza Española
Kilometers Cape Horn New
Zealand

! Figure 22.5 The voyage of HMS Beagle.

Darwin also spent much time thinking about geology. De- isms that enhance their survival and reproduction in specific
spite bouts of seasickness, he read Lyell’s Principles of Geology environments. Later, as he reassessed his observations, he be-
while aboard the Beagle. He experienced geologic change first- gan to perceive adaptation to the environment and the origin
hand when a violent earthquake rocked the coast of Chile, and of new species as closely related processes. Could a new species
he observed afterward that rocks along the coast had been arise from an ancestral form by the gradual accumulation of
thrust upward by several feet. Finding fossils of ocean organisms adaptations to a different environment? From studies made
high in the Andes, Darwin inferred that the rocks containing years after Darwin’s voyage, biologists have concluded that
the fossils must have been raised there by many similar earth- this is indeed what happened to the diverse group of Galápa-
quakes. These observations reinforced what he had learned gos finches (see Figure 1.22). The finches’ various beaks and
from Lyell: The physical evidence did not support the tradi- behaviors are adapted to the specific foods available on their
tional view that Earth was only a few thousand years old. home islands (Figure 22.6). Darwin realized that explaining
Darwin’s interest in the geographic distribution of species was such adaptations was essential to understanding evolution. As
further stimulated by the Beagle’s stop at the Galápagos, a group we’ll explore further, his explanation of how adaptations arise
of volcanic islands located near the equator about 900 km west centered on natural selection, a process in which individ-
of South America (Figure 22.5). Darwin was fascinated by the uals that have certain inherited traits tend to survive and
unusual organisms there. The birds he collected included the reproduce at higher rates than other individuals because of
finches mentioned in Chapter 1 and several kinds of mocking- those traits.
birds. These mockingbirds, though similar to each other, seemed By the early 1840s, Darwin had worked out the major fea-
to be different species. Some were unique to individual islands, tures of his hypothesis. He set these ideas on paper in 1844,
while others lived on two or more adjacent islands. Furthermore, when he wrote a long essay on descent with modification and
although the animals on the Galápagos resembled species living its underlying mechanism, natural selection. Yet he was still
on the South American mainland, most of the Galápagos species reluctant to publish his ideas, apparently because he antici-
were not known from anywhere else in the world. Darwin hy- pated the uproar they would cause. During this time, Darwin
pothesized that the Galápagos had been colonized by organisms continued to compile evidence in support of his hypothesis.
that had strayed from South America and then diversified, giv- By the mid-1850s, he had described his ideas to Lyell and a few
ing rise to new species on the various islands. others. Lyell, who was not yet convinced of evolution, never-
theless urged Darwin to publish on the subject before some-
one else came to the same conclusions and published first.
Darwin’s Focus on Adaptation
In June 1858, Lyell’s prediction came true. Darwin received
During the voyage of the Beagle, Darwin observed many ex- a manuscript from Alfred Russel Wallace (1823–1913), a British
amples of adaptations, inherited characteristics of organ- naturalist working in the South Pacific islands of the Malay

456 UNIT FOUR Mechanisms of Evolution

(a) Cactus-eater. The long, sharp beak of the (b) Insect-eater. The green warbler finch (c) Seed-eater. The large ground finch
cactus ground finch (Geospiza scandens) (Certhidea olivacea) uses its narrow, pointed (Geospiza magnirostris) has a large beak
helps it tear and eat cactus flowers and pulp. beak to grasp insects. adapted for cracking seeds that fall from
plants to the ground.

! Figure 22.6 Three examples of beak variation in Galápagos finches. The Galápagos Islands are
home to more than a dozen species of closely related finches, some found only on a single island. The most
striking differences among them are their beaks, which are adapted for specific diets.
MAKE CONNECTIONS Review Figure 1.22 (p. 17). To which of the other two species shown above is the
cactus-eater more closely related (that is, with which does it share a more recent common ancestor)?

Archipelago (see Figure 22.2). Wallace had developed a hy- “evolved”). Rather, he discussed descent with modification, a
pothesis of natural selection nearly identical to Darwin’s. He phrase that summarized his view of life. Organisms share
asked Darwin to evaluate his paper and forward it to Lyell if it many characteristics, leading Darwin to perceive unity in life.
merited publication. Darwin complied, writing to Lyell: “Your He attributed the unity of life to the descent of all organisms
words have come true with a vengeance. . . . I never saw a more from an ancestor that lived in the remote past. He also
striking coincidence . . . so all my originality, whatever it may thought that as the descendants of that ancestral organism
amount to, will be smashed.” On July 1, 1858, Lyell and a col- lived in various habitats over millions of years, they accumu-
league presented Wallace’s paper, along with extracts from lated diverse modifications, or adaptations, that fit them to
Darwin’s unpublished 1844 essay, to the Linnean Society of specific ways of life. Darwin reasoned that over long periods
London. Darwin quickly finished his book, titled On the Origin of time, descent with modification eventually led to the rich
of Species by Means of Natural Selection (commonly referred to as diversity of life we see today.
The Origin of Species), and published it the next year. Although Darwin viewed the history of life as a tree, with multiple
Wallace had submitted his ideas for publication first, he ad- branchings from a common trunk out to the tips of the
mired Darwin and thought that Darwin had developed the youngest twigs (Figure 22.7). The tips of the twigs represent
idea of natural selection so extensively that he should be the diversity of organisms living in the present. Each fork of
known as its main architect. the tree represents the most recent common ancestor of all the
Within a decade, Darwin’s book and its proponents had lines of evolution that subsequently branch from that point.
convinced most scientists that life’s diversity is the product of As an example, consider the three living species of elephants:
evolution. Darwin succeeded where previous evolutionists the Asian elephant (Elephas maximus) and African elephants
had failed, mainly by presenting a plausible scientific mecha-
nism with immaculate logic and an avalanche of evidence. " Figure 22.7 “I
think. . .” In this
1837 sketch, Darwin
The Origin of Species envisioned the
branching pattern
In his book, Darwin amassed evidence that descent with mod- of evolution.
ification by natural selection explains the three broad obser-
vations about nature listed in the Overview: the unity of life,
the diversity of life, and the match between organisms and
their environments.

Descent with Modification

In the first edition of The Origin of Species, Darwin never used
the word evolution (although the final word of the book is

CHAPTER 22 Descent with Modification: A Darwinian View of Life 457

(Loxodonta africana and L. cyclotis). These closely related species 22.8, fossils of extinct species can document the divergence of
are very similar because they shared the same line of descent present-day groups by “filling in” gaps between them.
until a relatively recent split from their common ancestor, as In his efforts at classification, Linnaeus had realized that
shown in the tree diagram in Figure 22.8. Note that seven lin- some organisms resemble each other more closely than others,
eages related to elephants have become extinct over the past but he had not linked these resemblances to evolution.
32 million years. As a result, there are no living species that fill Nonetheless, because he had recognized that the great diversity
the gap between the elephants and their nearest relatives to- of organisms could be organized into “groups subordinate to
day, the hyraxes and manatees. Such extinctions are not un- groups” (Darwin’s phrase), Linnaeus’s system meshed well with
common. In fact, many evolutionary branches, even some Darwin’s hypothesis. To Darwin, the Linnaean hierarchy re-
major ones, are dead ends: Scientists estimate that over 99% of flected the branching history of life, with organisms at the vari-
all species that have ever lived are now extinct. As in Figure ous levels related through descent from common ancestors.

Artificial Selection, Natural

Selection, and Adaptation
Hyracoidea Darwin proposed the mechanism of
(Hyraxes)
natural selection to explain the observ-
able patterns of evolution. He crafted
Sirenia
(Manatees his argument carefully, to persuade
and relatives) even the most skeptical readers. First he
†Moeritherium discussed familiar examples of selective
breeding of domesticated plants and
†Barytherium animals. Humans have modified other
species over many generations by se-
lecting and breeding individuals that
†Deinotherium possess desired traits, a process called
artificial selection (Figure 22.9). As a
result of artificial selection, crops, live-
†Mammut
stock animals, and pets often bear little
resemblance to their wild ancestors.
†Platybelodon Darwin then argued that a similar
process occurs in nature. He based his ar-
†Stegodon gument on two observations, from which
he drew two inferences:

†Mammuthus Observation #1: Members of a popu-

lation often vary in their inherited traits
(Figure 22.10).
Elephas maximus Observation #2: All species can pro-
(Asia)
duce more offspring than their environ-
ment can support (Figure 22.11), and
Loxodonta
africana many of these offspring fail to survive
(Africa) and reproduce.
Loxodonta cyclotis Inference #1: Individuals whose in-
(Africa)
herited traits give them a higher proba-
bility of surviving and reproducing in a
60 34 24 5.5 2 104 0
given environment tend to leave more
Millions of years ago Years ago offspring than other individuals.
Inference #2: This unequal ability of
! Figure 22.8 Descent with modification. This evolutionary tree of elephants and their individuals to survive and reproduce will
relatives is based mainly on fossils—their anatomy, order of appearance in strata, and geographic lead to the accumulation of favorable
distribution. Note that most branches of descent ended in extinction (denoted by the dagger traits in the population over generations.
symbol †). (Time line not to scale.)
Based on the tree shown here, approximately when did the most recent ancestor shared by Darwin saw an important connec-
? Mammuthus (woolly mammoths), Asian elephants, and African elephants live? tion between natural selection and the

458 UNIT FOUR Mechanisms of Evolution

# Figure 22.9 Artificial selection. These
different vegetables have all been selected from
one species of wild mustard. By selecting
variations in different parts of the plant,
breeders have obtained these divergent results.

Cabbage

Selection for
apical (tip) bud

Brussels sprouts Selection for Broccoli

Selection
axillary (side)
for flowers
buds
and stems

Selection
for stems

Selection
for leaves

Kale Wild mustard Kohlrabi

capacity of organisms to “overreproduce.” He began to make

this connection after reading an essay by economist Thomas
Malthus, who contended that much of human suffering—
disease, famine, and war—was the inescapable consequence
of the human population’s potential to increase faster than
food supplies and other resources. Darwin realized that the
capacity to overreproduce was characteristic of all species. Of
the many eggs laid, young born, and seeds spread, only a tiny
fraction complete their development and leave offspring of
their own. The rest are eaten, starved, diseased, unmated, or
unable to tolerate physical conditions of the environment
such as salinity or temperature.
! Figure 22.10 Variation in a population. Individuals in this
population of Asian ladybird beetles vary in color and spot pattern.
An organism’s heritable traits can influence not only its
Natural selection may act on these variations only if (1) they are own performance, but also how well its offspring cope with en-
heritable and (2) they affect the beetles’ ability to survive and vironmental challenges. For example, an organism might have
reproduce.
a trait that gives its offspring an advantage in escaping preda-
tors, obtaining food, or tolerating physical conditions. When
# Figure 22.11
Overproduction such advantages increase the number of offspring that survive
of offspring. A and reproduce, the traits that are favored will likely appear at
single puffball a greater frequency in the next generation. Thus, over time,
fungus can produce Spore
billions of offspring. cloud natural selection resulting from factors such as predators, lack
If all of these of food, or adverse physical conditions can lead to an increase
offspring and their in the proportion of favorable traits in a population.
descendants
How rapidly do such changes occur? Darwin reasoned that
survived to maturity,
they would carpet if artificial selection can bring about dramatic change in a rel-
the surrounding land atively short period of time, then natural selection should be
surface. capable of substantial modification of species over many hun-
dreds of generations. Even if the advantages of some heritable
traits over others are slight, the advantageous variations will
gradually accumulate in the population, and less favorable
variations will diminish. Over time, this process will increase
the frequency of individuals with favorable adaptations and
hence refine the match between organisms and their environ-
ment (see Figure 1.20).

CHAPTER 22 Descent with Modification: A Darwinian View of Life 459

Natural Selection: A Summary places or times. Natural selection is always operating, but
which traits are favored depends on the context in which a
Let’s now recap the main ideas of natural selection:
species lives and mates.
• Natural selection is a process in which individuals that Next, we’ll survey the wide range of observations that sup-
have certain heritable traits survive and reproduce at a port a Darwinian view of evolution by natural selection.
higher rate than other individuals because of those traits.
• Over time, natural selection can increase the match be- CONCEPT CHECK 22.2
tween organisms and their environment (Figure 22.12). 1. How does the concept of descent with modification
• If an environment changes, or if individuals move to a new explain both the unity and diversity of life?
environment, natural selection may result in adaptation to 2. WHAT IF? If you discovered a fossil of an extinct
these new conditions, sometimes giving rise to new species. mammal that lived high in the Andes, would you
One subtle but important point is that although natural se- predict that it would more closely resemble present-
lection occurs through interactions between individual organ- day mammals from South American jungles or
isms and their environment, individuals do not evolve. Rather, present-day mammals that live high in African
it is the population that evolves over time. mountains? Explain.
A second key point is that natural selection can amplify or 3. MAKE CONNECTIONS Review Figures 14.4 and 14.6
diminish only those heritable traits that differ among the in- (pp. 265 and 267) on the relationship between geno-
dividuals in a population. Thus, even if a trait is heritable, if type and phenotype. In a particular pea population,
all the individuals in a population are genetically identical for suppose that flowers with the white phenotype are fa-
that trait, evolution by natural selection cannot occur. vored by natural selection. Predict what would hap-
Third, remember that environmental factors vary from pen over time to the frequency of the p allele in the
place to place and over time. A trait that is favorable in one population, and explain your reasoning.
place or time may be useless—or even detrimental—in other For suggested answers, see Appendix A.

(a) A flower mantid

in Malaysia CONCEPT
22.3
Evolution is supported by
an overwhelming amount
of scientific evidence
In The Origin of Species, Darwin marshaled a broad range of ev-
idence to support the concept of descent with modification.
Still—as he readily acknowledged—there were instances in
which key evidence was lacking. For example, Darwin referred
to the origin of flowering plants as an “abominable mystery,”
(b) A leaf mantid and he lamented the lack of fossils showing how earlier
in Borneo
groups of organisms gave rise to new groups.
In the last 150 years, new discoveries have filled many of the
gaps that Darwin identified. The origin of flowering plants, for
example, is much better understood (see Chapter 30), and
many fossils have been discovered that signify the origin of
new groups of organisms (see Chapter 25). In this section,
we’ll consider four types of data that document the pattern of
evolution and illuminate the processes by which it occurs: di-
rect observations of evolution, homology, the fossil record,
and biogeography.
! Figure 22.12 Camouflage as an example of
evolutionary adaptation. Related species of the insects called
mantids have diverse shapes and colors that evolved in different Direct Observations of Evolutionary Change
environments.
Biologists have documented evolutionary change in thou-
Explain how these mantids demonstrate the three key observations
? about life introduced in this chapter’s Overview: the match between sands of scientific studies. We’ll examine many such studies
organisms and their environments, unity, and diversity. throughout this unit, but let’s look at two examples here.

460 UNIT FOUR Mechanisms of Evolution

Natural Selection in Response to Introduced Plant Species $ Figure 22.13 INQUIRY
Animals that eat plants, called herbivores, often have adaptations Can a change in a population’s food source
that help them feed efficiently on their primary food sources. result in evolution by natural selection?
What happens when herbivores begin to feed on a plant species
FIELD STUDY Soapberry bugs
with different characteristics than their usual food source? (Jadera haematoloma) feed
An opportunity to study this question in nature is provided most effectively when the
by soapberry bugs, which use their “beak,” a hollow, needle- length of their “beak” closely
matches the depth within the
like mouthpart, to feed on seeds located within the fruits of fruits of the seeds they eat. Scott
various plants. In southern Florida, the soapberry bug Jadera Carroll and his colleagues mea-
haematoloma feeds on the seeds of a native plant, the balloon sured beak lengths in soapberry
bug populations in southern
vine (Cardiospermum corindum). In central Florida, however,
Florida feeding on the native
balloon vines have become rare. Instead, soapberry bugs in balloon vine. They also mea-
that region now feed on goldenrain tree (Koelreuteria elegans), sured beak lengths in popula-
a species recently introduced from Asia. tions in central Florida feeding
on the introduced goldenrain
Soapberry bugs feed most effectively when their beak length tree, which has a flatter fruit
closely matches the depth at which the seeds are found within shape than the balloon vine. The
the fruit. Goldenrain tree fruit consists of three flat lobes, and researchers then compared the Soapberry bug with beak
measurements to those of mu- inserted in balloon vine fruit
its seeds are much closer to the fruit surface than the seeds of
seum specimens collected in the
the plump, round native balloon vine fruit. Researchers at the two areas before the goldenrain
University of Utah predicted that in populations that feed on tree was introduced.
goldenrain tree, natural selection would result in beaks that are RESULTS Beak lengths were shorter in populations feeding on the intro-
shorter than those in populations that feed on balloon vine duced species than in populations feeding on the native species, in which
the seeds are buried more deeply. The average beak length in museum
(Figure 22.13). Indeed, beak lengths are shorter in the popula-
specimens from each population (indicated by red
tions that feed on goldenrain tree. arrows) was similar to beak lengths in Beak
Researchers have also studied beak length evolution in populations feeding on native species.
soapberry bug populations that feed on plants introduced to
Louisiana, Oklahoma, and Australia. In each of these locations, 10 On native species,
the fruit of the introduced plants is larger than the fruit of the southern Florida
8
native plant. Thus, in populations feeding on introduced
species in these regions, the researchers predicted that natural 6

selection would result in the evolution of longer beak length. 4

Number of individuals

Again, data collected in field studies upheld this prediction. 2

The adaptation observed in these soapberry bug populations 0
had important consequences: In Australia, for example, the in-
Museum-specimen average
crease in beak length nearly doubled the success with which
soapberry bugs could eat the seeds of the introduced species. 10
Furthermore, since historical data show that the goldenrain 8 On introduced species,
central Florida
tree reached central Florida just 35 years before the scientific 6
studies were initiated, the results demonstrate that natural se- 4
lection can cause rapid evolution in a wild population.
2

The Evolution of Drug-Resistant Bacteria 0

6 7 8 9 10 11
An example of ongoing natural selection that dramatically Beak length (mm)
affects humans is the evolution of drug-resistant pathogens
CONCLUSION Museum specimens and contemporary data suggest
(disease-causing organisms and viruses). This is a particular that a change in the size of the soapberry bug’s food source can result
problem with bacteria and viruses because resistant strains of in evolution by natural selection for matching beak size.
these pathogens can proliferate very quickly. SOURCE S. P. Carroll and C. Boyd, Host race radiation in the soapberry
Consider the evolution of drug resistance in the bacterium bug: natural history with the history, Evolution 46: 1052–1069 (1992).
Staphylococcus aureus. About one in three people harbor this WHAT IF? When soapberry bug eggs from a population fed on bal-
species on their skin or in their nasal passages with no nega- loon vine fruits were reared on goldenrain tree fruits (or vice versa), the
tive effects. However, certain genetic varieties (strains) of this beak lengths of the adult insects matched those in the population from
which the eggs were obtained. Interpret these results.
species, known as methicillin-resistant S. aureus (MRSA), are

CHAPTER 22 Descent with Modification: A Darwinian View of Life 461

formidable pathogens. The past decade has seen an alarming $ Figure 22.14
increase in virulent forms of MRSA such as clone USA300, a I M PA C T
strain that can cause “flesh-eating disease” and potentially fa-
tal infections (Figure 22.14). How did clone USA300 and
other strains of MRSA become so dangerous? The Rise of MRSA
The story begins in 1943, when penicillin became the first
widely used antibiotic. Since then, penicillin and other antibi-
otics have saved millions of lives. However, by 1945, more
M ost methicillin-resistant Staphylococcus aureus (MRSA) infections
are caused by recently appearing strains such as clone USA300.
Resistant to multiple antibiotics and highly contagious, this strain
than 20% of the S. aureus strains seen in hospitals were already and its close relatives can cause lethal infections of the skin, lungs,
and blood. Researchers have identified key areas of the USA300
resistant to penicillin. These bacteria had an enzyme, penicil-
genome that code for its particularly virulent properties.
linase, that could destroy penicillin. Researchers responded by
The circular chromosome The highlighted regions
developing antibiotics that were not destroyed by penicilli- of clone USA300 has been contain genes that
nase, but some S. aureus populations developed resistance to sequenced and contains increase the strain’s
each new drug within a few years. 2,872,769 base pairs of DNA. virulence (see the key).

In 1959, doctors began using the powerful antibiotic

methicillin, but within two years, methicillin-resistant strains
1
of S. aureus appeared. How did these resistant strains emerge? 2,750,000
250,000 base pairs
Methicillin works by deactivating a protein that bacteria use
to synthesize their cell walls. However, S. aureus populations 2,500,000
exhibited variations in how strongly their members were af- Chromosome map
fected by the drug. In particular, some individuals were able to of S. aureus clone USA300 500,000

synthesize their cell walls using a different protein that was

2,250,000 Key to adaptations
not affected by methicillin. These individuals survived the
methicillin treatments and reproduced at higher rates than Methicillin resistance
Ability to colonize hosts 750,000
did other individuals. Over time, these resistant individuals
became increasingly common, leading to the spread of MRSA. Increased disease severity
Initially, MRSA could be controlled by antibiotics that
2,000,000 Increased gene exchange
(within species) and
worked differently from methicillin. But this has become toxin production 1,000,000
increasingly difficult because some MRSA strains are resistant
to multiple antibiotics—probably because bacteria can ex- 1,750,000
change genes with members of their own and other species 1,250,000
1,500,000
(see Figure 27.13). Thus, the present-day multidrug-resistant
strains may have emerged over time as MRSA strains that were WHY IT MATTERS MRSA infections have proliferated dramatically in the
past few decades, and the annual death toll in the United States is in the
resistant to different antibiotics exchanged genes. tens of thousands. There is grave concern about the continuing evolu-
The soapberry bug and S. aureus examples highlight two tion of drug resistance and the resulting difficulty of treating MRSA
key points about natural selection. First, natural selection is a infections. Ongoing studies of how MRSA strains colonize their hosts
process of editing, not a creative mechanism. A drug does not and cause disease may help scientists develop drugs to combat MRSA.
create resistant pathogens; it selects for resistant individuals FURTHER READING General information about MRSA can be found
that are already present in the population. Second, natural se- on the Centers for Disease Control and Prevention (CDC) website
(www.cdc.gov/mrsa) and in G. Taubes, The bacteria fight back,
lection depends on time and place. It favors those character-
Science 321:356–361 (2008).
istics in a genetically variable population that provide
WHAT IF? Efforts are underway to develop drugs that target S. au-
advantage in the current, local environment. What is benefi-
reus specifically and to develop drugs that slow the growth of MRSA
cial in one situation may be useless or even harmful in an- but do not kill it. Based on how natural selection works and on the
other. Beak lengths arise that match the size of the typical fruit fact that bacterial species can exchange genes, explain why each of
eaten by a particular soapberry bug population. However, a these strategies might be effective.
beak length suitable for fruit of one size can be disadvanta-
geous when the bug is feeding on fruit of another size. selection) in its descendants over time as they face different en-
vironmental conditions. As a result, related species can have
Homology characteristics that have an underlying similarity yet function
A second type of evidence for evolution comes from analyzing differently. Similarity resulting from common ancestry is
similarities among different organisms. As we’ve discussed, known as homology. As this section will explain, an under-
evolution is a process of descent with modification: Character- standing of homology can be used to make testable predictions
istics present in an ancestral organism are altered (by natural and explain observations that are otherwise puzzling.

462 UNIT FOUR Mechanisms of Evolution

 " Figure 22.15 Mammalian
forelimbs: homologous
structures. Even though they
have become adapted for different
functions, the forelimbs of all
Humerus mammals are constructed from the
same basic skeletal elements: one
large bone (purple), attached to
two smaller bones (orange and tan),
attached to several small bones
(gold), attached to several
Radius
metacarpals (green), attached to
Ulna approximately five digits, each of
which is composed of phalanges
Carpals (blue).
Metacarpals
Phalanges

Human Cat Whale Bat

Anatomical and Molecular Homologies ial structures if snakes and blind cave fishes had origins sepa-
rate from other vertebrate animals.
The view of evolution as a remodeling process leads to the pre-
Biologists also observe similarities among organisms at the
diction that closely related species should share similar fea-
molecular level. All forms of life use the same genetic language
tures—and they do. Of course, closely related species share the
of DNA and RNA, and the genetic code is essentially universal.
features used to determine their relationship, but they also
Thus, it is likely that all species descended from common ances-
share many other features. Some of these shared features make
tors that used this code. But molecular homologies go beyond a
little sense except in the context of evolution. For example, the
shared code. For example, organisms as dissimilar as humans
forelimbs of all mammals, including humans, cats, whales, and
and bacteria share genes inherited from a very distant common
bats, show the same arrangement of bones from the shoulder
ancestor. Some of these homologous genes have acquired new
to the tips of the digits, even though these appendages have
functions, while others, such as those coding for the ribosomal
very different functions: lifting, walking, swimming, and flying
subunits used in protein synthesis (see Figure 17.17), have re-
(Figure 22.15). Such striking anatomical resemblances would
tained their original functions. It is also common for organisms
be highly unlikely if these structures had arisen anew in each
to have genes that have lost their function, even though the ho-
species. Rather, the underlying skeletons of the arms, forelegs,
mologous genes in related species may be fully functional. Like
flippers, and wings of different mammals are homologous
vestigial structures, it appears that such inactive “pseudogenes”
structures that represent variations on a structural theme that
may be present simply because a common ancestor had them.
was present in their common ancestor.
Comparing early stages of development in different animal
species reveals additional anatomical homologies not visible
in adult organisms. For example, at some point in their devel-
opment, all vertebrate embryos have a tail located posterior to
Pharyngeal
(behind) the anus, as well as structures called pharyngeal pouches
(throat) pouches (Figure 22.16). These homologous throat
pouches ultimately develop into structures with very different
functions, such as gills in fishes and parts of the ears and
throat in humans and other mammals. Post-anal
tail
Some of the most intriguing homologies concern “left-
over” structures of marginal, if any, importance to the organ-
ism. These vestigial structures are remnants of features
Chick embryo (LM) Human embryo
that served a function in the organism’s ancestors. For in-
stance, the skeletons of some snakes retain vestiges of the ! Figure 22.16 Anatomical similarities in vertebrate
embryos. At some stage in their embryonic development, all
pelvis and leg bones of walking ancestors. Another example is
vertebrates have a tail located posterior to the anus (referred to as a
provided by eye remnants that are buried under scales in blind post-anal tail), as well as pharyngeal (throat) pouches. Descent from a
species of cave fishes. We would not expect to see these vestig- common ancestor can explain such similarities.

CHAPTER 22 Descent with Modification: A Darwinian View of Life 463

Homologies and “Tree Thinking” scended from ancestor 1 , whereas mammals, lizards and
Some homologous characteristics, such as the genetic code, snakes, crocodiles, and birds all descended from ancestor 3 . As
are shared by all species because they date to the deep ances- expected, the three homologies shown on the tree—limbs with
tral past. In contrast, homologous characteristics that evolved digits, the amnion (a protective embryonic membrane), and
more recently are shared only within smaller groups of organ- feathers—form a nested pattern. Limbs with digits were present
isms. Consider the tetrapods (from the Greek tetra, four, and in common ancestor 2 and hence are found in all of the de-
pod, foot), the vertebrate group that consists of amphibians, scendants of that ancestor (the tetrapods). The amnion was pres-
mammals, and reptiles (including birds—see Figure 22.17). All ent only in ancestor 3 and hence is shared only by some
tetrapods have limbs with digits (see Figure 22.15), whereas tetrapods (mammals and reptiles). Feathers were present only in
other vertebrates do not. Thus, homologous characteristics common ancestor 6 and hence are found only in birds.
form a nested pattern: All life shares the deepest layer, and To explore “tree thinking” further, note that in Figure 22.17,
each successive smaller group adds its own homologies to mammals are positioned closer to amphibians than to birds. As
those it shares with larger groups. This nested pattern is ex- a result, you might conclude that mammals are more closely re-
actly what we would expect to result from descent with mod- lated to amphibians than they are to birds. However, mammals
ification from a common ancestor. are actually more closely related to birds than to amphibians
Biologists often represent the pattern of descent from com- because mammals and birds share a more recent common an-
mon ancestors and the resulting homologies with an cestor (ancestor 3 ) than do mammals and amphibians (ances-
evolutionary tree, a diagram that reflects evolutionary rela- tor 2 ). Ancestor 2 is also the most recent common ancestor
tionships among groups of organisms. We will explore in de- of birds and amphibians, making mammals and birds equally
tail how evolutionary trees are constructed in Chapter 26, but related to amphibians. Finally, note that the tree in Figure 22.17
for now, let’s consider how we can interpret and use such trees. shows the relative timing of evolutionary events but not their
Figure 22.17 is an evolutionary tree of tetrapods and their actual dates. Thus, we can conclude that ancestor 2 lived be-
closest living relatives, the lungfishes. In this diagram, each fore ancestor 3 , but we do not know when that was.
branch point represents the common ancestor of all species that Evolutionary trees are hypotheses that summarize our cur-
descended from it. For example, lungfishes and all tetrapods de- rent understanding of patterns of descent. Our confidence in
these relationships, as with any hypothesis, depends on the
strength of the supporting data. In the case of Figure 22.17,
Each branch point represents the tree is supported by a variety of independent data
the common ancestor of
the lineages beginning sets, including both anatomical and DNA sequence
there and to the right of it. data. As a result, biologists feel confident that it
Lungfishes accurately reflects evolutionary history. As you
will read in Chapter 26, scientists can use
such well-supported evolutionary trees
1 Amphibians
Tetrapods

to make specific and sometimes surpris-

ing predictions about organisms.
Mammals
Amniotes

2
Digit-bearing A Different Cause of Resemblance:
limbs Convergent Evolution
3 Lizards
Amnion and snakes Although organisms that are closely
related share characteristics because
4 Crocodiles of common descent, distantly related
A hatch mark represents a
organisms can resemble one another
homologous characteristic
shared by all the groups 5 for a different reason: convergent
Ostriches
Birds

to the right of the mark. evolution, the independent evolu-

6 tion of similar features in different lin-
Feathers
Hawks and eages. Consider marsupial mammals,
other birds many of which live in Australia. Mar-
! Figure 22.17 Tree thinking: information provided in an evolutionary tree. supials are distinct from another group
This evolutionary tree for tetrapods and their closest living relatives, the lungfishes, is based on of mammals—the eutherians—few of
anatomical and DNA sequence data. The purple bars indicate the origin of three important
homologies, each of which evolved only once. Birds are nested within and evolved from reptiles; which live in Australia. (Eutherians
hence, the group of organisms called “reptiles” technically includes birds. complete their embryonic develop-
Are crocodiles more closely related to lizards or birds? Explain your answer. ment in the uterus, whereas marsupials
?

464 UNIT FOUR Mechanisms of Evolution

 gests that the reduction in the size of the pelvic bone may
have been driven by natural selection.
Fossils can also shed light on the origins of new groups of
NORTH organisms. An example is the fossil record of cetaceans, the
Sugar AMERICA
glider mammalian order that includes whales, dolphins, and por-
poises. Some of these fossils provided an unexpected line of
support for a hypothesis based on DNA data: that cetaceans
are closely related to even-toed ungulates, a group that in-
AUSTRALIA cludes deer, pigs, camels, and cows (Figure 22.19). What else
can fossils tell us about cetacean origins? The earliest
Flying cetaceans lived 50–60 million years ago. The fossil record in-
squirrel dicates that prior to that time, most mammals were terrestrial.
Although scientists had long realized that whales and other
! Figure 22.18 Convergent evolution. The ability to glide
through the air evolved independently in these two distantly related cetaceans originated from land mammals, few fossils had
mammals. been found that revealed how cetacean limb structure had
changed over time, leading eventually to the loss of hind
limbs and the development of flippers and tail flukes. In the
are born as embryos and complete their development in an past few decades, however, a series of remarkable fossils have
external pouch.) Some Australian marsupials have euther- been discovered in Pakistan, Egypt, and North America. These
ian look-alikes with superficially similar adaptations. For in- fossils document steps in the transition from life on land to
stance, a forest-dwelling Australian marsupial called the life in the sea, filling in some of the gaps between ancestral
sugar glider is superficially very similar to flying squirrels, and living cetaceans (Figure 22.20, on the next page).
gliding eutherians that live in North American forests Collectively, the recent fossil
(Figure 22.18). But the sugar glider has many other charac- discoveries document the forma-
teristics that make it a marsupial, much more closely related tion of new species and the origin
to kangaroos and other Australian marsupials than to flying of a major new group of mam-
squirrels or other eutherians. Once again, our understand- mals, the cetaceans. These discov-
ing of evolution can explain these observations. Although eries also show that cetaceans and ! Diacodexis, an early
they evolved independently from different ancestors, these their close living relatives (hip- even-toed ungulate
two mammals have adapted to similar environments in sim- popotamuses, pigs, deer, and
ilar ways. In such examples in which species share features other even-toed ungulates) are much more different from each
because of convergent evolution, the resemblance is said to other than were Pakicetus and early even-toed ungulates, such
be analogous, not homologous. Analogous features share as Diacodexis. Similar patterns are seen in fossils documenting
similar function, but not common ancestry, while homolo- the origins of other major new groups of organisms, including
gous features share common ancestry,
but not necessarily similar function.
Most mammals Cetaceans and even-toed ungulates
The Fossil Record
A third type of evidence for evolution
comes from fossils. As Chapter 25 dis-
cusses in more detail, the fossil record
documents the pattern of evolution,
showing that past organisms differed
from present-day organisms and that
many species have become extinct. Fos-
sils also show the evolutionary changes
(a) Canis (dog) (b) Pakicetus (c) Sus (pig) (d) Odocoileus (deer)
that have occurred in various groups of
organisms. To give one of hundreds of ! Figure 22.19 Ankle bones: one piece of the puzzle. Comparing fossils and present-
possible examples, researchers found day examples of the astragalus (a type of ankle bone) provides one line of evidence that cetaceans
that the pelvic bone in fossil stickleback are closely related to even-toed ungulates. (a) In most mammals, the astragalus is shaped like that
of a dog, with a double hump on one end (indicated by the red arrows) but not at the opposite
fish became greatly reduced in size over
end (blue arrow). (b) Fossils show that the early cetacean Pakicetus had an astragalus with double
time in a number of different lakes. The humps at both ends, a unique shape that is otherwise found only in even-toed ungulates, as
consistent nature of this change sug- shown here for (c) a pig and (d) a deer.

CHAPTER 22 Descent with Modification: A Darwinian View of Life 465

 Other
even-toed
ungulates

Hippopotamuses

†Pakicetus

†Rodhocetus
Common
m
ancestor
s
of cetaceans
t
†Dorudon

Living
cetaceans

70 60 50 40 30 20 10 0 Key to cetacean pelvis Pelvis Tibia

Millions of years ago and hind limb bones Femur Foot

! Figure 22.20 The transition to life in bones of extinct cetacean ancestors, including Which happened first during the evolution
the sea. Multiple lines of evidence support Pakicetus, Rodhocetus, and Dorudon. DNA ? of cetaceans: changes in hind limb structure
the hypothesis that cetaceans evolved from sequence data support the hypothesis that or the origin of tail flukes?
terrestrial mammals. Fossils document the cetaceans are most closely related to
reduction over time in the pelvis and hind limb hippopotamuses, even-toed ungulates.

mammals (see Chapter 25), flowering plants (see Chapter 30), horse species originated 5 million years ago in North America.
and tetrapods (see Chapter 34). In each of these cases, the fossil At that time, North and South America were close to their pres-
record shows that over time, descent with modification pro- ent locations, but they were not yet connected, making it diffi-
duced increasingly large differences among related groups of or- cult for horses to travel between them. Thus, we would predict
ganisms, ultimately resulting in the diversity of life we see today. that the oldest horse fossils should be found only on the conti-
nent on which horses originated—North America. This predic-
Biogeography tion and others like it for different groups of organisms have
A fourth type of evidence for evolution comes from been upheld, providing more evidence for evolution.
biogeography, the geographic distribution of species. The We can also use our understanding of evolution to explain
geographic distribution of organisms is influenced by many biogeographic data. For example, islands generally have many
factors, including continental drift, the slow movement of species of plants and animals that are endemic, which means
Earth’s continents over time. About 250 million years ago, they are found nowhere else in the world. Yet, as Darwin de-
these movements united all of Earth’s landmasses into a sin- scribed in The Origin of Species, most island species are closely re-
gle large continent called Pangaea (see Figure 25.14). lated to species from the nearest mainland or a neighboring
Roughly 200 million years ago, Pangaea began to break apart; island. He explained this observation by suggesting that islands
by 20 million years ago, the continents we know today were are colonized by species from the nearest mainland. These
within a few hundred kilometers of their present locations. colonists eventually give rise to new species as they adapt to
We can use our understanding of evolution and continental their new environments. Such a process also explains why two
drift to predict where fossils of different groups of organisms islands with similar environments in distant parts of the world
might be found. For example, scientists have constructed evolu- tend to be populated not by species that are closely related to
tionary trees for horses based on anatomical data. These trees each other, but rather by species related to those of the nearest
and the ages of fossils of horse ancestors suggest that present-day mainland, where the environment is often quite different.

466 UNIT FOUR Mechanisms of Evolution

What Is Theoretical About Darwin’s View of Life? years or less; see Chapter 24). Furthermore, as we’ll explore
throughout this unit, evolutionary biologists now recognize
Some people dismiss Darwin’s ideas as “just a theory.” However, that natural selection is not the only mechanism responsible
as we have seen, the pattern of evolution—the observation that for evolution. Indeed, the study of evolution today is livelier
life has evolved over time—has been documented directly and than ever as scientists find more ways to test predictions based
is supported by a great deal of evidence. In addition, Darwin’s on natural selection and other evolutionary mechanisms.
explanation of the process of evolution—that natural selection is Although Darwin’s theory attributes the diversity of life to
the primary cause of the observed pattern of evolutionary natural processes, the diverse products of evolution neverthe-
change—makes sense of massive amounts of data. The effects of less remain elegant and inspiring. As Darwin wrote in the fi-
natural selection also can be observed and tested in nature. nal sentence of The Origin of Species, “There is grandeur in this
What, then, is theoretical about evolution? Keep in mind view of life . . . [in which] endless forms most beautiful and
that the scientific meaning of the term theory is very different most wonderful have been, and are being, evolved.”
from its meaning in everyday use. The colloquial use of the
word theory comes close to what scientists mean by a hypoth-
esis. In science, a theory is more comprehensive than a hy- CONCEPT CHECK 22.3
pothesis. A theory, such as the theory of evolution by natural 1. Explain how the following statement is inaccurate:
selection, accounts for many observations and explains and “Antibiotics have created drug resistance in MRSA.”
integrates a great variety of phenomena. Such a unifying the- 2. How does evolution account for (a) the similar mam-
ory does not become widely accepted unless its predictions malian forelimbs with different functions shown in
stand up to thorough and continual testing by experiment Figure 22.15 and (b) the similar lifestyle of the two
and additional observation (see Chapter 1). As the next three distantly related mammals shown in Figure 22.18?
chapters demonstrate, this has certainly been the case with 3. WHAT IF? The fossil record shows that dinosaurs
the theory of evolution by natural selection. originated 200–250 million years ago. Would you ex-
The skepticism of scientists as they continue to test theo- pect the geographic distribution of early dinosaur fos-
ries prevents these ideas from becoming dogma. For example, sils to be broad (on many continents) or narrow (on
although Darwin thought that evolution was a very slow one or a few continents only)? Explain.
process, we now know that this isn’t always true. New species
For suggested answers, see Appendix A.
can form in relatively short periods of time (a few thousand

22 CHAPTER REVIEW
through the accumulation of adaptations. He refined his the-
SUMMARY OF KEY CONCEPTS ory for many years and finally published it in 1859 after learn-
ing that Wallace had come to the same idea.
CONCEPT 22.1 • In The Origin of Species, Darwin proposed that evolution occurs
The Darwinian revolution challenged traditional views of a by natural selection.
young Earth inhabited by unchanging species (pp. 453–455)
Observations
• Darwin proposed that life’s diversity arose from ancestral species
through natural selection, a departure from prevailing views. Individuals in a population Organisms produce more
• In contrast to catastrophism (the principle that events in the vary in their heritable offspring than the
past occurred suddenly by mechanisms not operating today), characteristics. environment can support.
Hutton and Lyell thought that geologic change results from
mechanisms that operated in the past in the same manner as at
the present time (uniformitarianism).
Inferences
• Lamarck hypothesized that species evolve, but the underlying
mechanisms he proposed are not supported by evidence. Individuals that are well suited
Why was the age of Earth important for Darwin’s ideas about to their environment tend to leave more
? evolution? offspring than other individuals.
and
CONCEPT 22.2 Over time, favorable traits
Descent with modification by natural selection explains the accumulate in the population.
adaptations of organisms and the unity and diversity of life
(pp. 455–460)
Describe how overreproduction and heritable variation relate to
• Darwin’s experiences during the voyage of the Beagle gave rise ? evolution by natural selection.
to his idea that new species originate from ancestral forms

CHAPTER 22 Descent with Modification: A Darwinian View of Life 467

CONCEPT 22.3 a. Humans and bats evolved by natural selection, and whales
evolved by Lamarckian mechanisms.
Evolution is supported by an overwhelming amount b. Forelimb evolution was adaptive in people and bats, but
of scientific evidence (pp. 460–467) not in whales.
• Researchers have directly observed natural selection leading c. Natural selection in an aquatic environment resulted in
to adaptive evolution in many studies, including research significant changes to whale forelimb anatomy.
on soapberry bug populations and on MRSA. d. Genes mutate faster in whales than in humans or bats.
• Organisms share characteristics because of common descent e. Whales are not properly classified as mammals.
(homology) or because natural selection affects independently 5. DNA sequences in many human genes are very similar to the
evolving species in similar environments in similar ways sequences of corresponding genes in chimpanzees. The most
(convergent evolution). likely explanation for this result is that
• Fossils show that past organisms differed from living organisms, a. humans and chimpanzees share a relatively recent com-
that many species have become extinct, and that species have mon ancestor.
evolved over long periods of time; fossils also document the b. humans evolved from chimpanzees.
origin of major new groups of organisms. c. chimpanzees evolved from humans.
• Evolutionary theory can explain biogeographic patterns. d. convergent evolution led to the DNA similarities.
e. humans and chimpanzees are not closely related.
Summarize the different lines of evidence supporting the hypothe-
? sis that cetaceans descended from land mammals and are closely
Level 3: Synthesis/Evaluation
related to even-toed ungulates.
6. EVOLUTION CONNECTION
Explain why anatomical and molecular features often fit a
similar nested pattern. In addition, describe a process that can
TEST YOUR UNDERSTANDING
cause this not to be the case.
Level 1: Knowledge/Comprehension 7. SCIENTIFIC INQUIRY
DRAW IT Mosquitoes resistant to the pesticide DDT first
1. Which of the following is not an observation or inference on appeared in India in 1959, but now are found throughout the
which natural selection is based? world. (a) Graph the data in the table below. (b) Examining
a. There is heritable variation among individuals. the graph, hypothesize why the percentage of mosquitoes re-
b. Poorly adapted individuals never produce offspring. sistant to DDT rose rapidly. (c) Suggest an explanation for the
c. Species produce more offspring than the environment can global spread of DDT resistance.
support.
d. Individuals whose characteristics are best suited to the en- Month 0 8 12
vironment generally leave more offspring than those 4% 45% 77%
Mosquitoes Resistant* to DDT
whose characteristics are less well suited.
e. Only a fraction of an individual’s offspring may survive. Source: C. F. Curtis et al., Selection for and against insecticide resistance and possible methods
of inhibiting the evolution of resistance in mosquitoes, Ecological Entomology 3:273–287 (1978).
2. Which of the following observations helped Darwin shape his *Mosquitoes were considered resistant if they were not killed within 1 hour of receiving a
concept of descent with modification? dose of 4% DDT.
a. Species diversity declines farther from the equator.
b. Fewer species live on islands than on the nearest continents. 8. WRITE ABOUT A THEME
c. Birds can be found on islands located farther from the main- Environmental Interactions Write a short essay (about
land than the birds’ maximum nonstop flight distance. 100–150 words) evaluating whether changes to an organ-
d. South American temperate plants are more similar to the ism’s physical environment are likely to result in evolution-
tropical plants of South America than to the temperate ary change. Use an example to support your reasoning.
plants of Europe.
e. Earthquakes reshape life by causing mass extinctions. For selected answers, see Appendix A.

Level 2: Application/Analysis
3. Within six months of effectively using methicillin to treat www.masteringbiology.com
S. aureus infections in a community, all new infections were
®
caused by MRSA. How can this result best be explained? 1. MasteringBiology Assignments:
a. S. aureus can resist vaccines. Tutorial Evidence for Evolution
b. A patient must have become infected with MRSA from an- Activities Artificial Selection • Darwin and the Galápagos Islands •
other community. The Voyage of the Beagle: Darwin’s Trip Around the World • Discovery
c. In response to the drug, S. aureus began making drug- Channel Video: Charles Darwin • Natural Selection for Antibiotic
resistant versions of the protein targeted by the drug. Resistance • Reconstructing Forelimbs
d. Some drug-resistant bacteria were present at the start of Questions Student Misconceptions • Reading Quiz • Multiple Choice •
treatment, and natural selection increased their frequency. End-of-Chapter
e. The drug caused the S. aureus DNA to change. 2. eText
4. The upper forelimbs of humans and bats have fairly similar Read your book online, search, take notes, highlight text, and more.
skeletal structures, whereas the corresponding bones in 3. The Study Area
whales have very different shapes and proportions. However, Practice Tests • Cumulative Test • 3-D Animations
genetic data suggest that all three kinds of organisms diverged • MP3 Tutor Sessions • Videos • Activities • Investigations • Lab
from a common ancestor at about the same time. Which of Media • Audio Glossary • Word Study Tools • Art
the following is the most likely explanation for these data?

468 UNIT FOUR Mechanisms of Evolution