4

Evolution: History and
Evidence

Charles Darwin described organic evolution as “descent with modification.”

Many of his ideas on how organic evolution happens were formulated dur-
ing and after his visit to the Galapagos Islands—the home of this land iguana
(Conolophus subcristatus).
Chapter Outline
4.1 Pre-Darwinian Theories of Change
Questions of the earth’s origin and life’s origin have been on the minds of humans 4.2 Darwin’s Early Years and His Journey
since prehistoric times, when accounts of creation were passed orally from generation Voyage of the HMS Beagle
4.3 Early Development of Darwin’s Ideas
to generation. For many people, these questions centered around concepts of purpose. of Evolution
Religious and philosophical writings help provide answers to such questions as: Why Geology
are we here? What is human nature really like? How do we deal with our mortality? Fossil Evidence
Galápagos Islands
Many of us are also concerned with other, very different, questions of origin: 4.4 The Theory of Evolution by Natural
How old is the planet earth? How long has life been on the earth? How did life arise Selection
on the earth? How did a certain animal species come into existence? Answers for Natural Selection
Adaptation
these questions come from a different authority—that of scientific inquiry. Alfred Russel Wallace
This chapter presents the history of the study of organic evolution and introduces 4.5 Microevolution, Macroevolution,
the theory of evolution by natural selection. Organic evolution, according to Charles and Evidence of Macroevolutionary
Change
­Darwin, is “descent with modification.” This simply means that populations change Biogeography
over time. Populations consist of individuals of the same species that occupy a given Paleontology
area at the same time. They share a unique set of genes. Population concepts are dis- Analogy and Homology
Interpreting the Evidence: Phylogeny
cussed in chapter 5. Evolution by itself does not imply any particular lineage or any and Common Descent
particular mechanism, and virtually all scientists agree that the evidence for change in
organisms over long time periods is overwhelming. Further, most scientists agree that
natural selection, the mechanism for evolution that Charles Darwin outlined, is one
explanation of how evolution occurs. In spite of the scientific certainty of evolution
and an acceptance of a general mechanism, much is still to be learned about the details
of evolutionary processes. Scientists will be debating these details for years to come.

4.1 P RE-D ARWINIAN T HEORIES OF C HANGE

LEARNING OUTCOME
1. Describe scientific thought on evolutionary change prior to the work of
Charles Darwin.

The idea of evolution did not originate with Charles Darwin. Some of the earliest
references to evolutionary change are from the ancient Greeks. The philosophers
­Empedocles (495–435 B.C.) and Aristotle (384–322 B.C.) described concepts of change
in living organisms over time. Georges-Louis Buffon (1707–1788) spent many years
studying comparative anatomy. His observations of structural variations in particular
organs of related animals convinced him that change must have occurred during the
history of life on earth. Buffon attributed change in organisms to the action of the
environment. He believed in a special creation of species and considered change
as being degenerate—for example, he described apes as degenerate ­humans. Eras-
mus ­Darwin (1731–1802), a physician and the grandfather of Charles Darwin, was
60 CHAPTER FOUR

intensely interested in questions of origin and change. He

­believed in the common ancestry of all organisms.
4.2 D ARWIN’S E ARLY Y EARS
Jean Baptiste Lamarck (1744–1829) was a distinguished AND H IS J OURNEY
French zoologist. His contributions to zoology include impor-
tant studies of animal classification. Lamarck published a set of LEARNING OUTCOME
invertebrate zoology books. His theory was based on a widely 1. Describe the circumstances that led to Charles Darwin
accepted theory of inheritance that organisms develop new ­becoming a naturalist on HMS Beagle.
organs, or modify existing organs, as needs arise. (Charles
Darwin also accepted this idea of inheritance.) Similarly, he Charles Robert Darwin (1809–1882) was born on February 12,
believed that disuse resulted in the degeneration of organs. 1809. His father, like his grandfather, was a physician. During
Lamarck believed that “need” was dictated by environmental Darwin’s youth in Shrewsbury, England, his interests centered
change and that change involved movement toward perfec- around dogs, collecting, and hunting birds—all popular
tion. The idea that change in a species is directed by need pastimes in wealthy families of nineteenth-century England.
logically led Lamarck to the conclusion that species could not These activities captivated him far more than the traditional
become extinct—they simply evolved into different species. education he received at boarding school. At the age of 16
Lamarck illustrated his ideas of change with the often- (1825), he entered medical school in Edinburgh, Scotland.
quoted example of the giraffe. He contended that ances- For two years, he enjoyed the company of the school’s well-
tral giraffes had short necks, much like those of any other established scientists. Darwin, however, was not interested
mammal. Straining to reach higher branches during brows- in a career in medicine because he could not bear the sight
ing resulted in their acquiring higher shoulders and longer of people experiencing pain. This prompted his father to
necks. These modifications, produced in one generation, suggest that he train for the clergy in the Church of England.
were passed on to the next generation. Lamarck published With this in mind, Charles enrolled at Christ’s College in
his theory in 1802 and included it in one of his invertebrate Cambridge and graduated with honors in 1831. This train-
zoology books, Philosophie Zoologique (1809). He defended ing, like the medical training he received, was disappointing
his ideas in spite of intense social criticism. for Darwin. Again, his most memorable experiences were
Lamarck’s acceptance of a theory of inheritance that those with Cambridge scientists. During his stay at Cam-
we now know is not correct led him to erroneous conclu- bridge, Darwin developed a keen interest in collecting bee-
sions about how evolution occurs. There is no evidence that tles and made valuable contributions to beetle taxonomy.
changes in the environment can initiate changes in organisms
that can be passed on to future generations. Instead, change Voyage of the HMS Beagle
originates in the process of gamete formation.
Random changes in the structure of DNA (mutation) and One of his Cambridge mentors, a botanist by the name of
chance processes involved in the assortment of genes into John S. Henslow, nominated Darwin to serve as a naturalist
gametes (e.g., independent assortment, crossing-over, and on a mapping expedition that was to travel around the world.
random fertilization—see chapter 3) result in variation among Darwin was commissioned as a naturalist on the HMS Beagle,
offspring. The environment then plays a role in determin- which set sail on December 27, 1831, on a five-year voyage
ing the survival of these variations in subsequent generations. (figure 4.1). Darwin helped with routine seafaring tasks and
Even though Larmarck’s mechanism of change was incorrect, made numerous collections, which he sent to Cambridge.
he should be remembered for his steadfastness in promoting The voyage gave him ample opportunity to explore tropical
the idea of evolutionary change and for his numerous accom- rain forests, fossil beds, the volcanic peaks of South America,
plishments in zoology. and the coral atolls of the South Pacific. Most important, Dar-
win spent five weeks on the Galápagos Islands, a group of
volcanic islands 900 km off the coast of Ecuador. Some of his
SECTION REVIEW 4.1 most revolutionary ideas came from his observations of plant
The idea of evolutionary change can be traced back to the and animal life on these islands. At the end of the voyage,
ancient Greek philosophers, and it persisted in the minds of Darwin was just 27 years old.
scientists until it was documented by Charles Darwin. Jean By 1842, Darwin had developed the essence of his con-
Baptist Lamarck believed that organisms undergo evolution- clusions but delayed their publication because of uncertainty
ary change in response to needs presented by environmental about how they would be received. His ideas were eventually
change. We now know that his ideas about how evolution presented before the Linnean Society in London in 1858, and
happens are wrong. On the Origin of Species by Means of Natural Selection was
­published in 1859 and revolutionized biology. In the years
Extinction is one possible outcome of evolutionary after his voyage, Darwin was an extremely prolific scientist.
change. If ­
Lamarck had been correct regarding the He published five volumes on Zoology of the Beagle Voyage
mechanism of evolutionary change, would extinction (1843), ­Fertilisation of Orchids (1862), The Variation of Plants
be more a more likely or a less likely outcome of evolu- and ­Animals under ­Domestication (1873), The Descent of
tionary change? Explain. Man (1871), and numerous other works.
 Evolution: History and Evidence 61

England
Asia
Europe
North Atlantic
America Ocean

Pacific
Cape
Ocean
Verde
Cocos Islands
Island Africa
`
Galapagos
Islands
Indian Ocean
South
America

Australia

Fernandina Isabela (Albemarle)

Santiago (James) Early Voyage from England

Santa Cruz (Indefatigable)

Return Voyage to England
San Cristobal (Chatham)

FIGURE 4.1 
Voyage of the HMS Beagle.  Charles Darwin grew up and was educated in England. He served as a naturalist on a five-year mapping expedition.
Darwin’s observations, especially those on the Galápagos Islands, were the basis for his theory of evolution by natural selection. The inset shows
two of the islands as photographed from the space shuttle Atlantis. Fernandina (bottom left of the insert) is composed of a single volcanic peak.
Isabela is composed of three volcanic peaks with the following names: Wolf (top left), Darwin (top center), and Alcedo (lower right).

SECTION REVIEW 4.2 The development of Darwin’s theory of evolution by natu-

ral selection was a long, painstaking process. Darwin had
The early years of life for Charles Darwin were spent collecting to become convinced that change occurs over time. Before
and hunting. These years promoted his keen interest in natu- leaving on his voyage, Darwin accepted the prevailing
ral history. While studying at Cambridge, he was nominated to opinion that the earth and its inhabitants had been created
serve as a naturalist on a world wide survey expedition on HMS 6,000  years ago and had not changed since. Because his
Beagle. His theory of evolution by natural selection was based observations during his voyage suggested that change does
on observations made during and after this voyage. occur, he realized that 6,000 years could not account for the
What were the three experiences in the life of Charles diversity of modern species if they arose through gradual
Darwin that were important in influencing the role he change. Once ideas of change were established in Darwin’s
played in helping us understand how evolution occurs? thinking, it took about 20 years of study to conceive, and
thoroughly document, the mechanism by which change
occurs. Darwin died without knowing the genetic principles
4.3 E ARLY D EVELOPMENT OF that support his theory.
D ARWIN’S I DEAS OF E VOLUTION
LEARNING OUTCOME Geology
1. List the sources of evidence that convinced Charles During his voyage, Darwin read Charles Lyell’s (1779–1875)
­Darwin that evolutionary change occurs. Principles of Geology. In this book, Lyell developed the
62 CHAPTER FOUR

ideas of another geologist, James Hutton, into the theory Galápagos Islands
of uniformitarianism. His theory was based on the idea
that the forces of wind, rain, rivers, volcanoes, and geo- On its trip up the western shore of South America, the HMS
logical uplift shape the earth today, just as they have in the Beagle stopped at the Galápagos Islands, which are named a­ fter
past. Lyell and Hutton contended that it was these forces, the large tortoises that inhabit them (Sp. galápago, ­tortoise).
not catastrophic events, that shaped the face of the earth The tortoises weigh up to 250 kg, have shells up to 1.8 m in
over hundreds of millions of years. This book planted two diameter, and live for 200 to 250 years. The islands’ governor
important ideas in Darwin’s mind: (1) the earth could be pointed out to Darwin that the shapes of the tortoise shells from
much older than 6,000 years and (2) if the face of the earth different parts of Albemarle Island differed. Darwin noticed
changed gradually over long periods, could not living forms other differences as well. Tortoises from the drier regions had
also change during that time? longer necks than tortoises from wetter habitats (figure 4.3). In
spite of their differences, the tortoises were quite similar to each
other and to the tortoises on the mainland of South America.
How could these overall similarities be explained?
Fossil Evidence Darwin reasoned that the island forms were derived from
Once the HMS Beagle reached South America, Darwin spent a few ancestral animals that managed to travel from the
time digging in the dry riverbeds of the pampas (grassy mainland, across 900 km of ocean. Because the Galápagos
plains) of Argentina. He found the fossil remains of an extinct Islands are volcanic (see figure 4.1) and arose out of the
hippopotamus-like animal, now called Toxodon, and fossils seabed, no land connection with the mainland ever existed.
of a horselike animal, Thoatherium. Both of these fossils One modern hypothesis is that tortoises floated from the
were from animals that were clearly different from any other mainland on mats of vegetation that regularly break free
animal living in the region. Modern horses were in South from coastal riverbanks during storms. Without predators
America, of course, but Spanish explorers had brought these on the islands, tortoises gradually increased in number.
horses to the Americas in the 1500s. The fossils suggested Darwin also explained some of the differences that
that horses had been present and had become extinct long he saw. In dryer regions, where vegetation was sparse, tor-
before the 1500s. Darwin also found fossils of giant armadil- toises with longer necks would be favored because they
los and giant sloths (figure 4.2). Except for their large size, could reach higher to get food. In moister regions, tortoises
these fossils were very similar to forms Darwin found living with longer necks would not necessarily be favored, and the
in the region. shorter-necked tortoises could survive.

   
(a) (b)

FIGURE 4.2 
The Giant Sloth. (a) Charles Darwin found evidence of the existence of giant sloths in South America similar to this Megatherium. Giant
sloths lived about 10,000 years ago and weighed in excess of 1,000 kg. They certainly did not move through tree branches like their only
living relative, Choloepus, 4.5 kg (b). Instead, they probably fed on leaves of lower tree branches that they could reach from the ground. The
similarity of giant sloths and modern-day sloths impressed Darwin with the fact that species change over time. Many species have become
extinct. As in this case, they often leave descendants that provide evidence of evolutionary change.
 Evolution: History and Evidence 63

(a)

FIGURE 4.3 
Galápagos Tortoises. (a) Shorter-necked subspecies of Chelonoidis
nigra* live in moister regions and feed on low-growing vegetation.
(b) Longer-necked subspecies live in drier regions and feed on
high-growing vegetation. This tortise, known as Lonesome George,
was the last of the G. nigra abingdonii subspecies. He died in
2012. *This species name replaces the older name (Geochelone
elephantopus) based on recent phylogenetic evidence. (b)

Darwin made similar observations of a group of dark, SECTION REVIEW 4.3

sparrow-like birds. Darwin noticed that the Galápagos finches
bore similarities suggestive of common ancestry. Scientists The evidence described by Charles Lyell convinced D ­ arwin
now think that Galápagos finches also descended from an that the earth was millions of years old, a time period long
ancestral species that originally inhabited the mainland of enough to account for changes in living organisms. South
South America. The chance arrival of a few finches, in either American fossils and observations of tortoises and finches
single or multiple colonization events, probably set up the on the Galápagos Islands convinced Darwin that change in
first bird populations on the islands. Early finches encoun- organisms over time does occur. These observations, and
tered many different habitats containing few other birds and others, were the basis for his description of the mechanism
predators. ­Ancestral finches, probably seed eaters, multiplied of evolutionary change.
rapidly and filled the seed-bearing habitats most attractive to How does the example of Galápagos finches illustrate
them. ­Fourteen species of finches arose from this ancestral the concept of adaptive radiation?
group, including one species found on small Cocos Island
northeast of the Galápagos Islands. Each species is adapted
to a specific habitat on the ­islands. The most obvious differ-
4.4 THE THEORY OF EVOLUTION
ence between these finches ­relates to dietary adaptations and BY NATURAL SELECTION
is reflected in the size and shape of their bills. The finches
of the Galápagos Islands provide an example of adaptive LEARNING OUTCOMES
radiation—the formation of new forms from an ancestral
1. Describe the four requirements for evolution to occur by
species, usually in response to the opening of new habitats
natural selection.
(figure 4.4).
2. Explain how reproductive success, phenotype, and envi-
Darwin’s experiences in South America and the Galápa-
ronment are related to evolutionary adaptation.
gos Islands convinced him that animals change over time. It
took the remaining years of his life for Darwin to formulate On his return to England in 1836, Darwin worked diligently
and document his ideas, and to publish a description of the on the notes and specimens he had collected and made
mechanism of evolutionary change. new observations. He was familiar with the obvious success
64 CHAPTER FOUR

FIGURE 4.4 
Adaptive Radiation of the Galápagos Finches.  Ancestral finches from the South American mainland colonized the Galápagos Islands.
Open habitats and few predators promoted the radiation of finches into 14 different species.

of breeders in developing desired variations in plant and ani- releases about 100,000 eggs with each spawning,
mal stocks (figure 4.5). He wondered if this artificial selec- a female sea star releases about 1 million eggs
tion of traits could have a parallel in the natural world. each season, and a female robin may lay four fertile
Ideas of how change occurred began to develop on his eggs each season. What if all of these eggs were
voyage. They took on their final form after 1838 when he read fertilized and developed to reproductive adults by the
an essay by Thomas Malthus (1766–1834) entitled Essay on the following year? A half million female sea stars (half
Principle of Population. Malthus believed that the human pop- of the million eggs would produce females and half
ulation has the potential to increase geometrically. (Geometric would produce males), each producing another
growth involves increasing by doubling or by some other mul- million eggs, repeated over just a few generations,
tiple rather than by adding a fixed number of individuals with would soon fill up the oceans! Even the adult female
each new generation.) However, because resources cannot keep robins, each producing four more robins, would
pace with the increased demands of a burgeoning population, result in unimaginable resource problems in just a
population-restraining factors, such as poverty, wars, plagues, few years.
and famine, begin to have an influence. Darwin realized that a 2. Inherited variations exist. They arise from a variety of
similar struggle to survive occurs in nature. When viewed over sources, including mutation, genetic recombination (see
generations, this struggle could be a means of natural ­selection. chapter 3), and random fertilization. Seldom are any two
Traits that were detrimental for an animal would be eliminated individuals exactly alike. Some of these genetic varia-
by the failure of the animal containing them to reproduce. tions may confer an advantage to the individual possess-
ing them. In other instances, variations may be harmful
to an individual. In still other instances, particular varia-
Natural Selection tions may be neither helpful nor harmful. (These are
said to be neutral.) These variations can be passed on to
Charles Darwin had no knowledge of modern genetic concepts,
offspring.
and therefore had no knowledge of the genetic principles that
3. Because resources are limited, existence is a constant
are the basis of evolutionary theory as it exists today. The mod-
struggle. Many more offspring are produced than
ern version of his theory can be summarized as follows:
resources can support; therefore, many individuals
1. All organisms have a far greater reproductive potential die. Darwin reasoned that the individuals that die are
than is ever realized. For example, a female oyster those with the traits ­(variations) that make survival and
 Evolution: History and Evidence 65

   
(a) (b) (c)

FIGURE 4.5 
Artificial Selection.  Dogs (Canis familiaris) were domesticated between 30,000 and 20,000 years ago. Although 99.9% genetically similar
to Canis lupis, the grey wolf (a), their ancestral wolf species is unknown. Since then, humans have been selectively breeding dogs for many
purposes. Some toy, or tea cup, dogs have primarily been bred for the enjoyment and status of the rich. Other dogs were bred for working.
The Shetland Sheep Dog (b) was bred for herding sheep in England. Still other dogs were bred for hunting. The Irish Wolfhound (c) originated
in Ireland and was used for hunting deer and wolves. Ancient Romans trained this breed to pull enemies from their horses during battle.

successful reproduction less likely. Traits that promote a variety of ways. Adaptations may be behavioral, physio-
successful reproduction are said to be adaptive. logical, or morphological. For example, arctic animals dis-
4. Adaptive traits become more common in subsequent play many adaptations to their environment. The arctic hare
­generations. Because organisms with maladaptive traits (Lepus arcticus) acquires a snow-white coat in the winter as
are less likely to reproduce, the maladaptive traits camouflage from predators. Its relatively small ears (as com-
become less frequent in a population. pared to other hares) help prevent heat loss as blood circu-
lates through peripheral vessels of its ears. Its huge feet (also
With these ideas, Darwin formulated a theory that in comparison to those of other hares) help keep the hare
explained how the tortoises and finches of the Galápagos on top of the snow as it moves around the arctic tundra. All
Islands changed over time. In addition, Darwin’s theory of these adaptations promote survival and make successful
explained how some animals, such as the ancient South ­reproduction more likely (figure 4.6).
American horses, could become extinct. What if a group of New adaptations arise as a result of mutations, and they
animals is faced with a new environment to which it is ill- are perpetuated by natural selection. Mutations are chance
adapted? Climatic changes, food shortages, and other envi- events, and most mutations are either harmful or neutral (see
ronmental stressors could lead to extinction. chapter 3). Adaptive mutations never occur as a result of a need,
and there is no guarantee that a species will change in order
to meet the challenges of a changing environment. If adaptive
Adaptation changes did occur in response to need, extinction would not
Adaptation occurs when a heritable change in a phenotype occur—and extinction is a fact of life for the majority of species.
increases an animal’s chances of successful reproduction
­ Most genetic variations exist as neutral alleles, having arisen
in a specified environment. Adaptations must be heritable by mutation years earlier, and are expressed as adaptive traits
changes to be passed to subsequent generations. Adaptations only when a population encounters a new environment and
are defined in the context of enhancing reproductive suc- natural selection acts on the population. Adaptation may result
cess because survival of a species occurs through success- in the evolution of multiple new groups if the environment can
ful reproduction, and survival of the species is the ultimate be exploited in different ways. When the evolution of multiple
measure of success. Adaptations are defined in the context of groups occurs, adaptive radiation results (see figure 4.4).
a specified environment because a change that promotes suc- Not every characteristic is an adaptation to some kind of
cessful reproduction in one environment may be detrimental environmental situation. An allele that provided some adap-
to reproductive success in a different environment. tive trait in one environment may be neutral when the envi-
Even though adaptations are defined in the context ronment changes, but persist in the population because the
of reproductive success, they can manifest themselves in trait is not detrimental. Other alleles may result in traits that
66 CHAPTER FOUR

judgments to evolution. On both of these matters, Darwin’s

ideas are closer to the views of most modern scientists.
Wallace’s work motivated Darwin to publish his own
ideas. The theory of natural selection, however, is usually
credited to Charles Darwin. Darwin’s years of work and mas-
sive accumulations of evidence led even Wallace to attribute
the theory to Darwin. Wallace wrote to Darwin in 1864:
I shall always maintain [the theory of evolution
by natural selection] to be actually yours and
yours only. You had worked it out in details I
had never thought of years before I had a ray of
light on the subject.

SECTION REVIEW 4.4

FIGURE 4.6 
Natural selection occurs because organisms have a high
Adaptations of the Arctic Hare (Lepus arcticus).  Lepus arcticus reproductive potential. Many variations within populations
lives in the arctic of Canada and Greenland on rocky slopes of
higher-elevation tundra, where it feeds on leaves, shoots, grasses, are inherited, and when organisms struggle for survival with
flowers, roots, and bark. Arctic hares show a variety of evolutionary other organisms or their environment, many individuals die.
adaptations for living in the arctic, including a snowy-white winter Those that survive pass adaptive traits, those traits that make
coat, relatively small ears to prevent excessive heat loss, and large them more likely to be successful at reproducing, on to their
feet that allow them to “snow-shoe” across the winter landscape. offspring. Maladaptive traits become less common, and thus
the population changes.
were never adaptive. Because these alleles result in neutral
traits, they are not selected against by natural selection and Is there a difference in thinking about natural selec-
persist in the population. tion as weeding out less fit variations versus selecting
for adaptive variations? Explain.
Alfred Russel Wallace
Alfred Russel Wallace (1823–1913) was an explorer of the 4.5 M ICROEVOLUTION,
­Amazon Valley and led a zoological expedition to the Malay
­Archipelago, which is an area of great biogeographical impor-
M ACROEVOLUTION,
tance. Wallace, like Darwin, was impressed with evolution- AND E VIDENCE OF
ary change and had read the writings of Thomas Malthus on
human populations. He synthesized a theory of evolution sim-
M ACROEVOLUTIONARY C HANGE
ilar to Darwin’s theory of evolution by natural selection. After
writing the details of his theory, Wallace sent his paper to Dar-
LEARNING OUTCOMES
win for criticism. Darwin recognized the similarity of Wallace’s 1. Compare microevolution and macroevolution.
ideas and prepared a short summary of his own theory. Both 2. Describe the sources of evidence for macroevolution.
Wallace’s and Darwin’s papers were published in the Journal
of the Proceedings of the Linnean Society in 1858. Darwin’s Organic evolution was defined earlier as a change in popula-
insistence on having Wallace’s ideas presented along with his tions over time, or simply “descent with modification.” The
own shows Darwin’s integrity. Darwin then shortened a man- change must involve the genetic makeup of the population
uscript he had been working on since 1856 and published it in order to be passed to future generations. These obser-
as On the Origin of Species by Means of Natural Selection in vations have led biologists to look for the mechanisms by
November 1859. The 1,250 copies prepared in the first print- which changes occur. There is no doubt that genetic changes
ing sold out the day the book was released. in populations occur—they have been directly observed in
In spite of the similarities in the theories of Wallace and the field and in the laboratory. These changes are the rea-
Darwin, there were also important differences. Wallace, for son that bacteria gain resistance to antibiotics and agricultural
example, believed that every evolutionary modification was a pests become resistant to pesticides. A change in the fre-
product of ­selection and, therefore, had to be adaptive for the quency of alleles in populations over time is called micro-
organism. Darwin, on the other hand, admitted that natural evolution. The processes that result in microevolution are
selection may not explain all evolutionary changes. He did not discussed in chapter 5.
insist on finding adaptive significance for every modification. Over longer timescales, microevolutionary processes
Further, unlike Darwin, Wallace stopped short of attributing result in large-scale changes. Large-scale changes that result
human intellectual functions and the ability to make moral in extinction and the formation of new species are called
 Evolution: History and Evidence 67

macroevolution. Macroevolutionary changes are difficult to Biogeography

observe in progress because of the geological timescales that
are usually involved. Evidence that macroevolution occurs, Biogeography is the study of the geographic distribution of
however, is compelling. This evidence is in the form of pat- plants and animals. Biogeographers try to explain why organ-
terns of plant and animal distribution, fossils, biochemical isms are distributed as they are. Biogeographic studies show
molecules, anatomical structures, and developmental pro- that life-forms in different parts of the world have distinctive
cesses. Just like a burglar cannot carry out a crime without evolutionary histories.
leaving some kind of evidence behind, organisms leave evi- One of the distribution patterns that biogeographers
dence of what they looked like and how they lived. Evolu- try to explain is how similar groups of organisms have
tionary investigators piece this evidence together and provide dispersed to places separated by seemingly impenetrable
detailed accounts of the lives of extinct organisms and their barriers. For example, native cats are inhabitants of most con-
relationships to modern forms. The sources of evidence for tinents of the earth, yet they cannot cross expanses of open
macroevolution are described in the next section. oceans. Obvious similarities suggest a common ancestry, but

(a) (b)

Bering Strait

Leopard (Panthera pardus)

Jaguar (Panthera onca)

(c)
FIGURE 4.7 
Biogeography as Evidence of Evolutionary Change. (a) The leopard (Panthera pardus) of Africa and Asia has a similar ecological role
to that of the (b) jaguar (Panthera onca) of Central and South America. Their similar form suggests common ancestry, even though they are
separated by apparently insurmountable oceanic barriers (c). Spotted varieties of these species are distinguished by the presence (jaguar) or
absence (leopard) of small spots within dark rosette markings of their coats. Biogeographers have provided probable explanations for these
observations. (See Evolutionary Insights, pages 75–76.)
68 CHAPTER FOUR

similarly obvious differences result from millions of years of uplift, in creating or removing barriers to the movements of
independent evolution (figure 4.7 and Evolutionary Insights, plants and animals. Biogeographers divide the world into six
pages 75–76). Biogeographers also try to explain why plants major biogeographic regions (figure 4.8). As they observe the
and animals, separated by geographical barriers, are often characteristic plants and animals in each of these regions
very different in spite of similar environments. For example, and learn about the earth’s geologic history, we understand
why are so many of the animals that inhabit Australia and more about animal distribution patterns and factors that
Tasmania so very different from animals in any other part played important roles in animal evolution. Only in under-
of the world? The major native herbivores of Australia and standing how the surface of the earth came to its present
Tasmania are the many species of kangaroos (Macropus). form can we understand its inhabitants.
In other parts of the world, members of the deer and cattle
groups fill these roles. Similarly, the Tasmanian wolf/tiger
(Thylacinus cynocephalus), now believed to be extinct,
Paleontology
was a predatory marsupial that was unlike any other large Paleontology (Gr. palaios, old 1 on, existing 1 logos, to
predator. Finally, biogeographers try to explain why oce- study), which is the study of the fossil record, provides some
anic islands often have relatively few, but unique, resident of the most direct evidence for evolution. Fossils (L. fossilis,
species. They try to document island colonization and sub- to dig) are evidence of plants and animals that existed in the
sequent evolutionary events, which may be very different past and have become incorporated into the earth’s crust
from the evolutionary events in ancestral, mainland groups. (e.g., as rock or mineral) (figure 4.9). Fossils are formed in
The discussion that follows will illustrate some of Charles sedimentary rock by a variety of methods. Most commonly,
Darwin’s conclusions about the island biogeography of the fossilization occurs when sediments (e.g., silt, sand, or vol-
Galápagos Islands. canic ash) quickly cover an organism to prevent scavenging
Modern evolutionary biologists recognize the impor- and in a way that seals out oxygen and slows decomposi-
tance of geological events, such as volcanic activity, the move- tion. As sediments continue to be piled on top of the dead
ment of great landmasses, climatic changes, and geological organism, pressures build. Water infiltrates the remains and

Palearctic region Nearctic

region

(a) (b)
Sahara Desert Oriental region Himalayan
Mountains (d)

Ethiopian
region Equator

Arabian (c)
Neotropical
Desert region

Australian
region

FIGURE 4.8 
Biogeographic Regions of the World.  Barriers, such as oceans, mountain ranges, and deserts, separate biogeographic regions of the world.
(a) The Sahara and Arabian Deserts separate the Ethiopian and Palearctic regions, (b) the Himalayan Mountains separate the Palearctic and
Oriental regions, (c) deep ocean channels separate the Oriental and Australian regions, and (d) the mountains of southern Mexico and
Mexico’s tropical lowlands separate the Nearctic and Neotropical regions.
 Evolution: History and Evidence 69

horses to be traced back about 55 million years (­figure 4.10).

Most of the evolutionary events occurred in what is now
North America. Hyracotherium was a dog-sized animal
(0.2  m in height at the shoulder). Fossils reveal the pres-
ence of four prominent toes on each foot and a tooth struc-
ture indicative of a browsing lifestyle. As the habitat became
more grassland-like, natural selection favored animals with
longer legs used for outrunning predators and larger more
durable teeth (molars) used for grazing. A loss of some foot
bones, and a reduction in others, was accompanied by an
elongation of the middle digit. The shift from browsing
to grazing was also accompanied by an elongation of the
face. Paleontologists have also used the fossil record to
describe the history of life on earth (see inside back cover).
Evidence from paleontology is clearly
Animation
some of the most convincing evidence Geological History
FIGURE 4.9 
of macroevolution. of the Earth
Paleontological Evidence of Evolutionary Change. Fossils,
such as these trilobites (Paradoxides), are direct evidence of
evolutionary change. Trilobites existed about 500 million years
ago and became extinct about 250 million years ago. Fossils may
form when an animal dies and is covered with sediments. Water
Analogy and Homology
dissolves calcium from hard body parts and replaces it with Structures and processes of organisms may be alike. There
another mineral, forming a hard replica of the original animal. This
process is called mineralization.
are two reasons for similarities, and both cases provide evi-
dence of evolution. Resemblance may occur when two unre-
lated organisms adapt to similar conditions. For example,
adaptations for flight have produced flat, gliding surfaces
in the wings of birds and insects. These similarities indicate
inorganic compounds and ions replace organic components. that independent evolution in these two groups produced
Hard parts of the organism are most likely to be fossilized, superficially similar structures and has allowed these two
but delicate structures are sometimes fossilized when silica groups of animals to exploit a common aerial environment.
is involved with replacement. These pressure and chemical The evolution of superficially similar structures in unrelated
changes transform the organism into a stony replica. Other organisms is called convergent evolution, and the simi-
fossilization processes form molds, casts of organisms, or lar structures are said to be analogous (Gr. analog 1 os,
carbon skeletons. Tracks and burrows, and even mummi- proportionate).
fied remains, are sometimes found. Fossilization is most Resemblance may also occur because two organisms
likely to occur in aquatic or semiaquatic environments. The share a common ancestry. Structures and processes in two
fossil record is therefore more complete for those groups of kinds of organisms that are derived from common ancestry
organisms living in or around water and for organisms with are said to be homologous (Gr. homolog 1 os, agreeing)
hard parts. (i.e., having the same or similar relation). Homology can
The fossil record provides information regarding involve aspects of an organism’s structure, and these homolo-
sequences in the appearance and disappearance of organ- gies are studied in the discipline called comparative anatomy.
isms. Different strata of rock result from differing rates Homology can also involve aspects of animal development
of sedimentation. Climatic and geological events influ- and function, and homologous processes are studied using
ence rates of sedimentation. When rates of sedimentation techniques of molecular biology.
change, a break in deposition occurs, leaving a distinct
layer or stratum. Successive strata are piled on top of each
other, with younger strata on top of older strata. Fossils in Comparative Anatomy
younger strata are of animals that lived more recently than Comparative anatomy is the study of the structure of liv-
fossils in older strata. Geologists can correlate strata around ing and fossilized animals and the homologies that indicate
the world and determine when strata were formed using evolutionarily close relationships. In many cases, homolo-
radioactive dating (How Do We Know Evolutionary Times- gies are readily apparent. For example, vertebrate append-
cales?, page 72). ages have a common arrangement of bones that can be
Paleontologists use this information to provide an traced back through primitive amphibians and certain
understanding of many evolutionary lineages. Many verte- groups of fish. Appendages have been modified for differ-
brate lineages are very well documented in the fossil ent functions such as swimming, running, flying, and grasp-
record. For example, the fossil record allows the history of ing, but the basic sets of bones and the relationships of
70 CHAPTER FOUR

Millions of
years before
present Equus (500 kg)
Holocene
Pleistocene Pliohippus
Pliocene
Hypohippus
10 Merychippus (100 kg)
Miocene

Parahippus

20 Miohippus
Oligocene

30 Mesohippus (45 kg)

Haplohippus

40 Epihippus

Orohippus
Eocene

Key

Primarily Grazer
50 Hyracotherium (20 kg)
Grazer/Browser

Primarily Browser

60

FIGURE 4.10 
Reconstruction of an Evolutionary Lineage from Evidence in the Fossil Record.  The fossil record allows horse evolution to be traced
back about 55 million years. The horse ancestors illustrated were not direct ancestor/descendant sequences. The illustrations depict anatomical
changes that occurred during horse evolution. Horse ancestors were small, primarily browsing animals that walked on the tips of 3 or 4 toes.
Evolution resulted in larger animals adapted to a grazing lifestyle and that walked or ran on the tips of their middle toe digits. Note that
evolutionary lineages are seldom simple ladders of change. Instead, numerous evolutionary side branches often meet with extinction.

bones to each other have been retained (figure 4.11). The eardrum (tympanic membrane) to the inner ear. Studies of
similarities in structure of these bones reflect their common the fossil record reveal the origin of these middle ear bones
ancestry and the fact that vertebrate appendages, although (­figure 4.12). Small bones involved in jaw suspension in prim-
modified in their details of structure, have retained their itive fish are incorporated into the remnants of a pharyngeal
primary functions of locomotion. (gill) slit to form the middle ear. In amphibians, reptiles, and
Other structures may be homologous even though they birds, a single bone of fish (the hyomandibular bone) forms
differ in appearance and function. The origin of the middle ear the middle ear bone (the columella or stapes). In the evolu-
bones of terrestrial vertebrates provides an example. Fish do tion of mammals, two additional bones that contributed to jaw
not have middle and outer ears. Their inner ear provides for support in ancient fish (the quadrate and articular bones) are
the senses of equilibrium, balance, and hearing, with sound used in the middle ear (the incus and malleus, respectively).
waves being transmitted through bones of the skull. Terres- There are many other examples of structures that have
trial vertebrates evolved from primitive fish, and the evolution changed from an ancestral form. Sometimes these changes
of life on land resulted in an ear that could detect airborne result in a structure that can be detrimental to the organism.
vibrations. These vibrations are transmitted to receptors of the The human vermiform appendix evolved from a large fermen-
inner ear through a middle ear and, in some cases, an outer tation pouch, and it is still used in this manner in animals like
ear. One (amphibians, reptiles, and birds) or more (mammals) rabbits and many other herbivores. In humans, the vermiform
small bones of the middle ear transmit vibrations from the appendix may have functions related to the lymphatic system,
 Evolution: History and Evidence 71

Horse 3
1 Quadrate
Articular
4 (a)
Columella (stapes)
Bat
5
Inner ear

Tympanic
Brain membrane
1 2
3 Middle
ear

Quadrate
3
Human Bird Articular
Throat
1

(b)
5 2
4
3

Brain
1

5 Lizard
4
3 Quadrate
4 Whale
1 2 Articular
3 2
Throat

FIGURE 4.11 
The Concept of Homology.  The forelimbs of vertebrates evolved
from an ancestral pattern. Even vertebrates as dissimilar as whales (c)
and bats have the same basic arrangement of bones. The digits Stapes
(fingers) are numbered 1 (thumb) to 5 (little finger). Color coding Incus
indicates homologous bones. Malleus

but these functions pale in comparison to the problems that

result when it becomes infected. Boa constrictors have min-
ute remnants of hindlimb (pelvic) bones that are remnants of
the appendages of their reptilian ancestors. Baleen whales,
like all whales, evolved from land mammals and also retain
remnants of the pelvic appendages their ancestors used for
walking on land. In both of these cases the remnant pelvic
appendages serve no apparent function (figure 4.13). Struc-
tures that have no apparent function in modern animals but Eustachian tube
clearly evolved from functioning structures in ancestors are Tympanic membrane
Outer ear
called vestigial structures and provide another source of (d)
evidence of evolutionary change.
FIGURE 4.12 
Evolution of the Vertebrate Ear Ossicles. (a) Lateral view of the
Developmental Patterns skull of a primitive amphibian showing the two bones (quadrate and
Evidence of evolution also comes from observing the devel- articular) that function in jaw support and contribute to the middle
opmental patterns of organisms. The developmental stages ear bones of mammals. Diagrammatic sections of the heads of
of related animals often retain common features because (b) a primitive amphibian, (c) a primitive reptile, and (d) a mammal
showing the fate of three bones of primitive fish. The columella
changes in the genes that control the development of ani- (stapes) is derived from a bone called the hyomandibular, the incus
mals are usually harmful and are eliminated by natural selec- is derived from the quadrate bone, and the malleus is derived from
tion. For example, early embryonic stages of vertebrates the articular bone.
72 CHAPTER FOUR

FIGURE 4.13 
Vestigial Structures.  The pelvic bones of baleen whales evolved from functional pelvic bones of the whales’ terrestrial mammalian
ancestor. These bones have no apparent function in whales and are an example of a vestigial structure.

How Do We Know Evolutionary Timescales?

E volutionary timescales have

been studied for many
years using relative dat-
ing techniques that place geologic
events in sequential order that is
The half-life of uranium 238 is 4.5
billion years. Since no radioactive
isotope is added to a rock once it
is formed, the ratio of radioactive
isotope to its decay product can be
selected against, so the “clock” for
changes in these molecules runs very
slowly. In other molecules, changes
are less detrimental to function and
are tolerated. The “clock” for change
determined by the geological record. used to date a rock, and a fossil in these molecules runs less slowly.
Absolute dating provides absolute contained within that rock. Because This dating principle also works in
dates for events and is determined different isotopes have different regions of DNA that do not code
by radiometric and molecular tech- rates of decay, varying time frames for functional proteins. If the rate
niques. These techniques have can be measured. The very long of change in a region of DNA is rela-
been used to provide the timescales half-life of uranium-238 makes it tively constant over time, the amount
shown in the table on the inside useful for dating rock formations of change can be used to date evolu-
back cover of this textbook. that are hundreds of millions of tionary events. This concept is called
Igneous rocks form when lava years old. Carbon-14 has a half-life the molecular clock. The accuracy
cools. These rocks contain radio- of 5,730 years and is used to date of the molecular clock is controver-
active isotopes of elements. For fossils thousands of years old. sial, but when dates can be substan-
example, uranium-238 is the radio- Molecular techniques have tiated by radiometric and relative
active isotope that decays through provided another absolute dating dating techniques, it is a very effec-
a series of isotopes to produce technique, the molecular clock. tive tool for evolutionary biologists.
lead-206. These decay processes Molecular biologists have found that The timescales depicted in the Evolu-
occur at a constant rate for a par- within each kind of molecule, the tionary Insights reading in this chapter
ticular isotope. The rate of decay is rate of change is relatively constant. are an example of how molecular
described in terms of the isotope’s Different molecules have different and radiometric techniques can be
half-life—the time required for one- rates of change. Changes in some used to establish when evolutionary
half of the unstable nuclei to decay. molecules are detrimental and events occurred.
 Evolution: History and Evidence 73

Molecular Biology
Within the last 25 years, molecular biology has yielded a
wealth of information on evolutionary relationships. Studying
changes in anatomical structures and physiological processes
Fish Bird Mammal
reflects genetic change and evolution. Unfortunately, it is
(a) (chicken) (human) often difficult to sort out the relationship between genes and
the structures and functions they control. Studying nuclear
DNA, mitochondrial DNA, ribosomal RNA, and proteins is
particularly useful in evolutionary biology because these mol-
Forebrain ecules can provide direct evidence of changes in genes and
thus evolution. Just as animals can have homologous struc-
Midbrain tures, animals also have homologous biochemical processes
that can be studied using molecular biological methods.
Molecular methods have several advantages: they are
Hindbrain
useful with all organisms, the data are quantifiable with read-
ily available computer software, and databases of molecular
information for many organisms are very large and growing.
The use of molecular data allows biologists to investigate the
causes of the genetic variation and molecular processes that
Neural influence evolution. These data also provide information for
tube
the construction of evolutionary trees (phylogenies).
The principle behind molecular analysis is that closely
related organisms will be genetically more similar than dis-
tantly related organisms. Genetic similarity or degree of relat-
edness is reflected in the variation (or lack of it) in the amino
acids that comprise a protein or in the bases that comprise
DNA. This genetic variation can be quantified in a number of
ways. Genetic variation is often measured by the proportion
Mammal Bird of polymorphic loci in a population. A polymorphic locus
(b) (human) (chicken)
is one where two or more alleles exist. For example, imagine
FIGURE 4.14  a researcher examined 20 loci from representatives of two
Developmental Patterns. (a) The early embryonic stages of populations. In the first population, the researcher found that
various vertebrates are remarkably similar. These similarities result five of these loci had more than one allele. The proportion of
in the preservation of developmental sequences that evolved in polymorphic loci would be 5∕20 or 0.25. In the second popu-
early common ancestors of vertebrates. (b) Organ systems, like the lation, 10 of these loci had more than one allele, and the pro-
nervous system, also show similar developmental patterns. Later
developmental differences may result from evolutionary changes in portion of polymorphic loci was 0.5. Genetic variation could
the timing of developmental events. be greater in the second population for a ­number of rea-
sons. For example, it could indicate greater time since diver-
gence from an ancestor and thus more time for variations to
are remarkably similar (figure 4.14a). Many organ systems accumulate, or it could indicate genetic mixing with more
of vertebrates also show similar developmental patterns than one ancestral group. Documenting genetic variation is
(­figure 4.14b). These similarities are compelling evidence of important in evolutionary studies because this variation is the
evolutionary relationships within animal groups. fuel for natural selection. (Recall that genetic variation was
Adults in vertebrate groups are obviously different from the second of four points in the earlier description of natural
one another. If developmental patterns are so similar, how did selection.)
differences in adult stages arise? Evolution is again the answer. Techniques for isolating and manipulating DNA have
These differences are a result of evolutionary changes in the provided very powerful tools for the analysis of genetic varia-
genes that control the onset of developmental stages and the tion among groups of animals. The polymerase chain reac-
rate at which development occurs. These changes result in dif- tion (PCR) and automated DNA sequencers allow researchers
ferences in the size and proportions of organs. Differing growth to begin with very small amounts of sample DNA and quickly
rates of the bones of the skull, for example, can explain the and inexpensively determine the base sequences of DNA
proportional differences ­between bones of the human and ape and other genetic fingerprinting patterns. Variation in DNA
skulls. Modern developmental biology is providing a growing in homologous genes and regions suggests relationships
appreciation of how evolution has conserved many genes that between genes and groups of organisms. The tree dia­gram
control the developmental similarities of animal groups. At the on the inside of the front cover of this textbook is based
same time, it is helping to explain developmental changes that largely on the study of variation in the base sequence of ribo-
result in the great diversity of animal life. somal RNA.
74 CHAPTER FOUR

How Do We Know about Evolution—“Evo-Devo”?

T he study of development
has revealed that animals
in groups as diverse as
insects and humans share genes that
direct certain stages of development.
are finding that the same Hox genes
direct the development of structures in
diverse groups of animals. Body seg-
mentation is present in both arthro-
pods (insects and their relatives) and
both groups. The study of evolution
through the analysis of development
is sometimes called “evo-devo” and
is revealing that a relatively small set
of common genes underlies basic
Homeobox (Hox) genes determine vertebrates. In the past, this observa- developmental processes in many
the identity of body regions in early tion has been explained as a case of organisms. Evo-devo is helping to
embryos. They identify, for example, convergent evolution. It now appears explain how small changes in these
where a limb of a fly or a fish will be that the same gene appears to regu- development-directing genes can
located. Mutations in these genes late the development of segmentation have far-reaching evolutionary con-
may cause body parts to appear in in both groups, which means that sequences. (See the Evolutionary
the wrong place, to be duplicated, or body segmentation was probably Insights in chapters 7 and 18 for other
to be lost. Developmental biologists present in a common ancestor of examples.)

Interpreting the Evidence: 600– 800

million years
Phylogeny and Common Descent ago

Scientists use the data gathered from studies just described

to understand how organisms are related to each other.
Phylogeny refers to the evolutionary relationships among 450– 500
species. It includes the depiction of ancestral species and the
relationships of modern descendants of a common ances-
tor. These depictions involve the use of phylogenetic trees Branch
showing lines of descent. In the past, biologists relied mainly >300 Branch point (node)
on the fossil record and anatomical traits in the construction ~260
of phylogenetic trees. The addition of molecular data is revo- 150– 200
lutionizing phylogenetic studies.
100–140
Phylogenies are reconstructed by examining variations
in homologous structures, proteins, and genes. The process 40–50 Pseudogene
~35 40–80
begins by gathering data from different organisms being com- linkage
pared. In molecular studies it usually involves sampling DNA
Myoglobin α1 ψα1 θ1 ζ ε Gγ Aγ δ β
or proteins from present-day organisms. Proteins or DNA
from a particular locus or set of loci are compared using com- α family β family
puter programs that include assumptions about the types of
changes that are more likely to occur. (Changes of purine FIGURE 4.15 
to pyrimidine, or vice versa, are less likely than changes of A Phylogenetic Tree of Hemoglobin.  This phylogenetic tree
one purine to another purine or one pyrimidine to another is derived from molecular studies of the hemoglobin molecule.
The numbers associated with each branch point indicate the
pyrimidine.) The computer program examines all possible
approximate time in millions of years. The pseudogene in the
relationships between the different organisms and groups alpha family is a gene that is apparently nonfunctional.
the organisms based on the fewest number of evolutionary
changes that must have taken place since they shared a com-
mon ancestor. points or nodes that represent ancestral molecules. In other
Figure 4.15 shows a phylogenetic tree for the hemoglo- trees (see Evolutionary Insights box figure 4.1), nodes could
bin molecule. Hemoglobin genes are grouped into alpha and represent individuals, populations, or species. A branch repre-
beta families. The products of these genes are incorporated into sents an evolutionary connection between molecules (individu-
the hemoglobin molecule that transports oxygen in red blood als, populations, or species). The longer the branch, the greater
cells. Another related gene codes for the oxygen storage pigment the variation and the more distant the evolutionary relationship
in muscle, myoglobin. Genes in the beta family have branch between molecules (individuals, populations, or species). The
 Evolution: History and Evidence 75

E VOLUTIONARY I NSIGHTS
An Example from Big-Cat Phylogeny

A ll sources of evidence are used in piecing together the

evolutionary relationships of animals. Evidence from
paleontology, biogeography, molecular biology, and
ecology is being used to understand the evolutionary past of big-
cat species.
Another closely related cat is the tiger (Panthera tigris), which
was historically found in Turkey, India, China, and Indonesia.
Fossils show that a tiger/jaguar-like cat existed in northern China
and may be the ancestor of these three species. One hypothesis
is that this common ancestor spread into Europe and also east-
Paleontologists have established that the earliest cats arose ward. About 1 million years ago it crossed the Bering land bridge
approximately 30 million years ago. The earliest known cat fossil that connected Asia and North America. The breakup of the Ber-
(Proailurus) is from a bobcat-sized animal that lived in Europe ing land bridge then isolated these groups. The jaguar’s current
about 25 million years ago. Paleontologists have uncovered distribution in Central and South America may be explained by
numerous lineages. All but one of these has met with extinction. the appearance of lions (Felix concolor) in North America. Com-
Modern cats apparently arose from a single common ancestor petition may have driven jaguars southward.
about 10 million years ago. Molecular phylogenetic studies have begun to sort out cat
Biogeographers have helped to explain the distribution of relationships that have been impossible to sort out using other
wild cats around the globe. Working from the fossil record and methods. These studies expanded a traditional interpretation of
a knowledge of current distribution patterns, biogeographers three major lineages that arose about 10 million years ago into
develop hypotheses to explain why certain cat species are dis- a total of eight smaller lineages. Molecular studies within the
tributed as they are. For example, the leopard (Panthera par- Panthera lineage suggest that species within this group origi-
dus) and the jaguar (Panthera onca) are remarkably similar in nated from a common ancestor between 3 and 6 million years
appearance (see figure 4.7) and ecological roles. In the wild, ago. This date is based on studies of base sequence differ-
their similar appearance presents no problem in the identifica- ences in certain ribosomal RNA genes and mitochondrial genes
tion of these species because of their distribution. The leopard among Panthera species. A larger percent difference (in this
is found in Africa, the Middle East, and Asia. The jaguar is found case about 7.5%) in base pairs of the same gene means a longer
only in Central America and northern and central South America. time elapsed since the common ancestor. Box figure 4.1 shows
Proailurus (extinct)

Pseudaelurus (extinct)
Extinct lineages including the
saber-toothed cats
Smilodon and Homotherium
Caracal group

Domestic cat group

Bay cat group

Puma group
Lynx group
Ocelot group

Asian leopard cat group

Panthera group

Jaguar

Tiger
Lion

Leopard
Snow leopard
Clouded leopard

BOX FIGURE 4.1  A Partial Cat Phylogeny.   Eight lineages of modern cats were derived from a single ancestor about 10 million
years ago. All cats trace their ancestry back about 30 million years. The inset shows one hypothesis for the relationships between
modern species of the Panthera lineage, which originated about 6 million years ago.
(Continued)
76 CHAPTER FOUR

one interpretation of the phylogeny of this lineage based on use powerful neck and limb muscles to cache prey high in the
the analysis of base pair differences in the NADH-5 mitochon- boughs of trees. Antelope and other prey may be three times the
drial gene. body weight of the leopard. This behavior reduces competition
Morphological (structural) and behavioral differences between from scavenging hyenas and opportunistic lions that may happen
leopards and jaguars may be explained by habitat differences. upon a leopard’s kill.
Most jaguars are found in the densely forested areas of the Ama- Both of these species are threatened by habitat loss and
zon Basin. Their smaller size is thought to be an adaptation to hunting—both have been prized for their fur. Although the leop-
climatic and vegetational changes encountered as the cats moved ard has a very large range and diverse prey base, a number of
south. Leopards, on the other hand, have evolved into a com- subspecies are gone from many parts of their original range. Jag-
plex group of subspecies as they adapted to diverse environments uars are severely threatened by deforestation. It is estimated that
across their range. One habit of African and Asian leopards is to there are 15,000 individuals left in the wild.

relatively short branch distance between nodes in the beta fam- SECTION REVIEW 4.5
ily reflects close evolutionary ties. Similar conclusions are drawn
for genes in the alpha family. The conclusions drawn from this Microevolution is a change in the frequency of alleles in
tree are that all of the modern hemoglobin genes are derived populations over time. Macroevolution involves large-scale
from a single gene that existed between 600 and 800 million changes such as the formation of new species. Evidence of
years ago. Phylogenetic trees like this one, and the many that macroevolutionary change is compelling. Sources of evi-
will follow in chapters 7 through 22, reinforce our ideas of com- dence include the following: biogeography, paleontology,
mon descent. All life is related, and the evidence of this rela- comparative anatomy, developmental biology, and molecu-
tionship is overwhelming. lar biology. All of these lines of evidence contribute to our
Evolution is the major unifying theme in biology understanding of phylogenetic relationships.
because it helps explain both the similarities and the diver-
sity of life. There is no doubt that it has occurred in the past Some opponents of evolutionary theory contend that
and continues to occur today. Chapter 5 examines how the evolutionary theory is not valid science because it
principles of population genetics have been combined with concerns events of the past that cannot be observed or
Darwinian evolutionary theory into what is often called the ­re-created in the laboratory. How would you respond
modern synthesis. to this criticism?

S UMMARY selection and Malthus’s theory of human population growth

helped shape his theory.
4.1 Pre-Darwinian Theories of Change 4.4 The Theory of Evolution by Natural Selection
Organic evolution is the change of a species over time. Darwin’s theory of natural selection includes the follow-
Ideas of evolutionary change can be traced back to the ing elements: (1) All organisms have a greater reproductive
ancient Greeks. potential than is ever attained; (2) inherited variations arise
Jean Baptiste Lamarck was an eighteenth-century proponent by mutation; (3) in a constant struggle for existence, those
of evolution and proposed a mechanism—inheritance of organisms that are least suited to their environment die; and
acquired characteristics—to explain it. (4) the adaptive traits present in the survivors tend to be
passed on to subsequent generations, and the nonadaptive
4.2 Darwin’s Early Years and His Journey traits tend to be lost.
Charles Darwin saw impressive evidence for evolutionary Adaptation may refer to a process of change or a result of
change while on a mapping expedition on the HMS Beagle. change. An adaptation is a characteristic that increases an
The theory of uniformitarianism, South American fossils, organism’s potential to reproduce in a given environment.
and observations of tortoises and finches on the Galápagos
Not all evolutionary changes are adaptive, nor do all evolu-
Islands convinced Darwin that evolution occurs.
tionary changes lead to perfect solutions to environmental
4.3 Early Development of Darwin’s Ideas of Evolution problems.
After returning from his voyage, Darwin began formulating Alfred Russel Wallace outlined a theory similar to Darwin’s
his theory of evolution by natural selection. In addition to but never accumulated significant evidence documenting his
his experiences on his voyage, later observations of artificial theory.
 Evolution: History and Evidence 77

4.5 Microevolution, Macroevolution, and Evidence of 4. Resemblance between two organisms can occur because of
Macroevolutionary Change shared ancestry. Structures that are similar because of shared
Microevolution is the change in the frequency of alleles in ancestry are said to be
populations over time. Macroevolution is large-scale change a. homologous.
that results in extinction and the formation of new species b. analogous.
over geological timescales. c. convergent.
Evidence of macroevolutionary change comes from the study d. adaptive.
of biogeography, paleontology, comparative anatomy, molec-
5. Millions of years ago two populations of a species were sepa-
ular biology, and developmental biology.
rated as a result of two continents drifting apart. Members
All sources of evidence are used in studying the phylogeny of
of these two populations adapted to different environments
animals. These studies have resulted in the wealth of informa-
and today are very different from one another and classified
tion on animal lineages that will be presented in chapters that
as different species. ­Accounts like this that explain how geo-
follow.
graphical barriers help ­account for the evolution of groups of
animals is the work of
a. paleontologists.
C ONCEPT R EVIEW Q UESTIONS b. molecular biologists.
1. Charles Lyell’s Principles of Geology influenced Charles Dar- c. comparative anatomists.
win’s ideas of evolutionary change in which of the following d. biogeographers.
ways?
a. His description of fossils of South America convinced
Darwin that species that were present in the past became A NALYSIS AND A PPLICATION
extinct, thus reinforcing ideas of change.
b. Lyell developed the ideas of another geologist, James Hutton, Q UESTIONS
who advocated that uniformitarianism, not catastrophic
1. Outline a hypothesis and design a test of “inheritance of
change, was responsible for shaping the face of the earth.
acquired characteristics,” and define what is meant by the
c. Lyell’s work convinced Darwin that the earth must be much word “theory” in the theory of evolution by natural selection.
older than 6,000 years.
2. Describe the implications of inheritance of acquired character-
d. Lyell described the volcanic origin of the Galápagos Islands, istics for our modern concepts of how genes function.
providing Darwin with ideas of how the islands might have
3. How would you explain the presence of gaps in the fossil
been colonized by tortoises and finches.
record? Would you be more likely to see gaps in the fossil
e. Both b and c are correct. record of rodents, fish, molluscs, or segmented worms? Explain
f. All of the above (a–d) are correct. your answer.
2. The evolution of a number of new characteristics from 4. Why is the stipulation of “a specified environment” included in
an ancestral form is called ____________ and results from the definition of adaptation?
____________. 5. Imagine that you could go back in time and meet simulta-
a. natural selection; the opening of new habitats neously with Charles Darwin and Gregor Mendel. Construct
b. adaptive radiation; the opening of new habitats a dialogue in which you explain to both the effect of their
c. punctuated equilibrium; loss of genetic variation in a ideas on each other’s theories and their theories on modern
species biology. Include their responses and questions throughout
the dialogue.
d. microevolution; extinction of one of two closely related
species
3. A heritable change that increases an animal’s chance of suc-
cessful reproduction
a. is an adaptation.
Enhance your study of this chapter with study tools and practice
b. prevents evolutionary change.
tests. Also ask your instructor about the resources available through
c. always results in the evolution of a new species. Connect, including a media-rich eBook, interactive learning tools,
d. is called genetic drift. and animations.