468 Chapter 18 | Evolution and the Origin of Species

18.1 | Understanding Evolution

By the end of this section, you will be able to:
• Describe how the present-day theory of evolution was developed
• Define adaptation
• Explain convergent and divergent evolution
• Describe homologous and vestigial structures
• Discuss misconceptions about the theory of evolution

Evolution by natural selection describes a mechanism for how species change over time. That species change had been
suggested and debated well before Darwin began to explore this idea. The view that species were static and unchanging was
grounded in the writings of Plato, yet there were also ancient Greeks who expressed evolutionary ideas. In the eighteenth
century, ideas about the evolution of animals were reintroduced by the naturalist Georges-Louis Leclerc Comte de Buffon
who observed that various geographic regions have different plant and animal populations, even when the environments are
similar. It was also accepted that there were extinct species.
During this time, James Hutton, a Scottish naturalist, proposed that geological change occurred gradually by the
accumulation of small changes from processes operating like they are today over long periods of time. This contrasted
with the predominant view that the geology of the planet was a consequence of catastrophic events occurring during a
relatively brief past. Hutton’s view was popularized in the nineteenth century by the geologist Charles Lyell who became
a friend to Darwin. Lyell’s ideas were influential on Darwin’s thinking: Lyell’s notion of the greater age of Earth gave
more time for gradual change in species, and the process of change provided an analogy for gradual change in species. In
the early nineteenth century, Jean-Baptiste Lamarck published a book that detailed a mechanism for evolutionary change.
This mechanism is now referred to as an inheritance of acquired characteristics by which modifications in an individual are
caused by its environment, or the use or disuse of a structure during its lifetime, could be inherited by its offspring and thus
bring about change in a species. While this mechanism for evolutionary change was discredited, Lamarck’s ideas were an
important influence on evolutionary thought.

Charles Darwin and Natural Selection

In the mid-nineteenth century, the actual mechanism for evolution was independently conceived of and described by two
naturalists: Charles Darwin and Alfred Russel Wallace. Importantly, each naturalist spent time exploring the natural world
on expeditions to the tropics. From 1831 to 1836, Darwin traveled around the world on H.M.S. Beagle, including stops
in South America, Australia, and the southern tip of Africa. Wallace traveled to Brazil to collect insects in the Amazon
rainforest from 1848 to 1852 and to the Malay Archipelago from 1854 to 1862. Darwin’s journey, like Wallace’s later
journeys to the Malay Archipelago, included stops at several island chains, the last being the Galápagos Islands west of
Ecuador. On these islands, Darwin observed species of organisms on different islands that were clearly similar, yet had
distinct differences. For example, the ground finches inhabiting the Galápagos Islands comprised several species with
a unique beak shape (Figure 18.2). The species on the islands had a graded series of beak sizes and shapes with very
small differences between the most similar. He observed that these finches closely resembled another finch species on the
mainland of South America. Darwin imagined that the island species might be species modified from one of the original
mainland species. Upon further study, he realized that the varied beaks of each finch helped the birds acquire a specific
type of food. For example, seed-eating finches had stronger, thicker beaks for breaking seeds, and insect-eating finches had
spear-like beaks for stabbing their prey.

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Chapter 18 | Evolution and the Origin of Species 469

Figure 18.2 Darwin observed that beak shape varies among finch species. He postulated that the beak of an ancestral
species had adapted over time to equip the finches to acquire different food sources.

Wallace and Darwin both observed similar patterns in other organisms and they independently developed the same
explanation for how and why such changes could take place. Darwin called this mechanism natural selection. Natural
selection, also known as “survival of the fittest,” is the more prolific reproduction of individuals with favorable traits that
survive environmental change because of those traits; this leads to evolutionary change.
For example, a population of giant tortoises found in the Galapagos Archipelago was observed by Darwin to have longer
necks than those that lived on other islands with dry lowlands. These tortoises were “selected” because they could reach
more leaves and access more food than those with short necks. In times of drought when fewer leaves would be available,
those that could reach more leaves had a better chance to eat and survive than those that couldn’t reach the food source.
Consequently, long-necked tortoises would be more likely to be reproductively successful and pass the long-necked trait to
their offspring. Over time, only long-necked tortoises would be present in the population.
Natural selection, Darwin argued, was an inevitable outcome of three principles that operated in nature. First, most
characteristics of organisms are inherited, or passed from parent to offspring. Although no one, including Darwin and
Wallace, knew how this happened at the time, it was a common understanding. Second, more offspring are produced than
are able to survive, so resources for survival and reproduction are limited. The capacity for reproduction in all organisms
outstrips the availability of resources to support their numbers. Thus, there is competition for those resources in each
generation. Both Darwin and Wallace’s understanding of this principle came from reading an essay by the economist
Thomas Malthus who discussed this principle in relation to human populations. Third, offspring vary among each other in
regard to their characteristics and those variations are inherited. Darwin and Wallace reasoned that offspring with inherited
characteristics which allow them to best compete for limited resources will survive and have more offspring than those
individuals with variations that are less able to compete. Because characteristics are inherited, these traits will be better
represented in the next generation. This will lead to change in populations over generations in a process that Darwin
called descent with modification. Ultimately, natural selection leads to greater adaptation of the population to its local
environment; it is the only mechanism known for adaptive evolution.
Papers by Darwin and Wallace (Figure 18.3) presenting the idea of natural selection were read together in 1858 before the
Linnean Society in London. The following year Darwin’s book, On the Origin of Species, was published. His book outlined
in considerable detail his arguments for evolution by natural selection.
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Figure 18.3 Both (a) Charles Darwin and (b) Alfred Wallace wrote scientific papers on natural selection that were
presented together before the Linnean Society in 1858.

Demonstrations of evolution by natural selection are time consuming and difficult to obtain. One of the best examples has
been demonstrated in the very birds that helped to inspire Darwin’s theory: the Galápagos finches. Peter and Rosemary
Grant and their colleagues have studied Galápagos finch populations every year since 1976 and have provided important
demonstrations of natural selection. The Grants found changes from one generation to the next in the distribution of beak
shapes with the medium ground finch on the Galápagos island of Daphne Major. The birds have inherited variation in the
bill shape with some birds having wide deep bills and others having thinner bills. During a period in which rainfall was
higher than normal because of an El Niño, the large hard seeds that large-billed birds ate were reduced in number; however,
there was an abundance of the small soft seeds which the small-billed birds ate. Therefore, survival and reproduction were
much better in the following years for the small-billed birds. In the years following this El Niño, the Grants measured beak
sizes in the population and found that the average bill size was smaller. Since bill size is an inherited trait, parents with
smaller bills had more offspring and the size of bills had evolved to be smaller. As conditions improved in 1987 and larger
seeds became more available, the trend toward smaller average bill size ceased.

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Chapter 18 | Evolution and the Origin of Species 471

Field Biologist
Many people hike, explore caves, scuba dive, or climb mountains for recreation. People often participate
in these activities hoping to see wildlife. Experiencing the outdoors can be incredibly enjoyable and
invigorating. What if your job was to be outside in the wilderness? Field biologists by definition work outdoors
in the “field.” The term field in this case refers to any location outdoors, even under water. A field biologist
typically focuses research on a certain species, group of organisms, or a single habitat (Figure 18.4).

Figure 18.4 A field biologist tranquilizes a polar bear for study. (credit: Karen Rhode)

One objective of many field biologists includes discovering new species that have never been recorded.
Not only do such findings expand our understanding of the natural world, but they also lead to important
innovations in fields such as medicine and agriculture. Plant and microbial species, in particular, can reveal
new medicinal and nutritive knowledge. Other organisms can play key roles in ecosystems or be considered
rare and in need of protection. When discovered, these important species can be used as evidence for
environmental regulations and laws.

Processes and Patterns of Evolution

Natural selection can only take place if there is variation, or differences, among individuals in a population. Importantly,
these differences must have some genetic basis; otherwise, the selection will not lead to change in the next generation.
This is critical because variation among individuals can be caused by non-genetic reasons such as an individual being taller
because of better nutrition rather than different genes.
Genetic diversity in a population comes from two main mechanisms: mutation and sexual reproduction. Mutation, a change
in DNA, is the ultimate source of new alleles, or new genetic variation in any population. The genetic changes caused by
mutation can have one of three outcomes on the phenotype. A mutation affects the phenotype of the organism in a way
that gives it reduced fitness—lower likelihood of survival or fewer offspring. A mutation may produce a phenotype with a
beneficial effect on fitness. And, many mutations will also have no effect on the fitness of the phenotype; these are called
neutral mutations. Mutations may also have a whole range of effect sizes on the fitness of the organism that expresses
them in their phenotype, from a small effect to a great effect. Sexual reproduction also leads to genetic diversity: when two
parents reproduce, unique combinations of alleles assemble to produce the unique genotypes and thus phenotypes in each
of the offspring.
A heritable trait that helps the survival and reproduction of an organism in its present environment is called an adaptation.
Scientists describe groups of organisms becoming adapted to their environment when a change in the range of genetic
variation occurs over time that increases or maintains the “fit” of the population to its environment. The webbed feet
of platypuses are an adaptation for swimming. The snow leopards’ thick fur is an adaptation for living in the cold. The
cheetahs’ fast speed is an adaptation for catching prey.
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Whether or not a trait is favorable depends on the environmental conditions at the time. The same traits are not always
selected because environmental conditions can change. For example, consider a species of plant that grew in a moist climate
and did not need to conserve water. Large leaves were selected because they allowed the plant to obtain more energy from
the sun. Large leaves require more water to maintain than small leaves, and the moist environment provided favorable
conditions to support large leaves. After thousands of years, the climate changed, and the area no longer had excess water.
The direction of natural selection shifted so that plants with small leaves were selected because those populations were able
to conserve water to survive the new environmental conditions.
The evolution of species has resulted in enormous variation in form and function. Sometimes, evolution gives rise to groups
of organisms that become tremendously different from each other. When two species evolve in diverse directions from a
common point, it is called divergent evolution. Such divergent evolution can be seen in the forms of the reproductive
organs of flowering plants which share the same basic anatomies; however, they can look very different as a result of
selection in different physical environments and adaptation to different kinds of pollinators (Figure 18.5).

Figure 18.5 Flowering plants evolved from a common ancestor. Notice that the (a) dense blazing star (Liatrus spicata)
and the (b) purple coneflower (Echinacea purpurea) vary in appearance, yet both share a similar basic morphology.
(credit a: modification of work by Drew Avery; credit b: modification of work by Cory Zanker)

In other cases, similar phenotypes evolve independently in distantly related species. For example, flight has evolved in both
bats and insects, and they both have structures we refer to as wings, which are adaptations to flight. However, the wings
of bats and insects have evolved from very different original structures. This phenomenon is called convergent evolution ,
where similar traits evolve independently in species that do not share a common ancestry. The two species came to the same
function, flying, but did so separately from each other.
These physical changes occur over enormous spans of time and help explain how evolution occurs. Natural selection acts on
individual organisms, which in turn can shape an entire species. Although natural selection may work in a single generation
on an individual, it can take thousands or even millions of years for the genotype of an entire species to evolve. It is over
these large time spans that life on earth has changed and continues to change.

Evidence of Evolution
The evidence for evolution is compelling and extensive. Looking at every level of organization in living systems, biologists
see the signature of past and present evolution. Darwin dedicated a large portion of his book, On the Origin of Species, to
identifying patterns in nature that were consistent with evolution, and since Darwin, our understanding has become clearer
and broader.
Fossils
Fossils provide solid evidence that organisms from the past are not the same as those found today, and fossils show a
progression of evolution. Scientists determine the age of fossils and categorize them from all over the world to determine
when the organisms lived relative to each other. The resulting fossil record tells the story of the past and shows the evolution
of form over millions of years (Figure 18.6). For example, scientists have recovered highly detailed records showing the
evolution of humans and horses (Figure 18.6). The whale flipper shares a similar morphology to appendages of birds and
mammals (Figure 18.7) indicating that these species share a common ancestor.

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Chapter 18 | Evolution and the Origin of Species 473

Figure 18.6 In this (a) display, fossil hominids are arranged from oldest (bottom) to newest (top). As hominids evolved,
the shape of the skull changed. An artist’s rendition of (b) extinct species of the genus Equus reveals that these ancient
species resembled the modern horse (Equus ferus) but varied in size.

Anatomy and Embryology

Another type of evidence for evolution is the presence of structures in organisms that share the same basic form. For
example, the bones in the appendages of a human, dog, bird, and whale all share the same overall construction (Figure 18.7)
resulting from their origin in the appendages of a common ancestor. Over time, evolution led to changes in the shapes and
sizes of these bones in different species, but they have maintained the same overall layout. Scientists call these synonymous
parts homologous structures.

Figure 18.7 The similar construction of these appendages indicates that these organisms share a common ancestor.

Some structures exist in organisms that have no apparent function at all, and appear to be residual parts from a past common
ancestor. These unused structures without function are called vestigial structures. Other examples of vestigial structures
are wings on flightless birds, leaves on some cacti, and hind leg bones in whales.

Visit this interactive site (http://openstaxcollege.org/l/bone_structures) to guess which bones structures are
homologous and which are analogous, and see examples of evolutionary adaptations to illustrate these concepts.

Another evidence of evolution is the convergence of form in organisms that share similar environments. For example,
species of unrelated animals, such as the arctic fox and ptarmigan, living in the arctic region have been selected for seasonal
474 Chapter 18 | Evolution and the Origin of Species

white phenotypes during winter to blend with the snow and ice (Figure 18.8ab). These similarities occur not because of
common ancestry, but because of similar selection pressures—the benefits of not being seen by predators.

Figure 18.8 The white winter coat of the (a) arctic fox and the (b) ptarmigan’s plumage are adaptations to their
environments. (credit a: modification of work by Keith Morehouse)

Embryology, the study of the development of the anatomy of an organism to its adult form, also provides evidence
of relatedness between now widely divergent groups of organisms. Mutational tweaking in the embryo can have such
magnified consequences in the adult that embryo formation tends to be conserved. As a result, structures that are absent
in some groups often appear in their embryonic forms and disappear by the time the adult or juvenile form is reached. For
example, all vertebrate embryos, including humans, exhibit gill slits and tails at some point in their early development.
These disappear in the adults of terrestrial groups but are maintained in adult forms of aquatic groups such as fish and some
amphibians. Great ape embryos, including humans, have a tail structure during their development that is lost by the time of
birth.
Biogeography
The geographic distribution of organisms on the planet follows patterns that are best explained by evolution in conjunction
with the movement of tectonic plates over geological time. Broad groups that evolved before the breakup of the
supercontinent Pangaea (about 200 million years ago) are distributed worldwide. Groups that evolved since the breakup
appear uniquely in regions of the planet, such as the unique flora and fauna of northern continents that formed from the
supercontinent Laurasia and of the southern continents that formed from the supercontinent Gondwana. The presence of
members of the plant family Proteaceae in Australia, southern Africa, and South America is best by their presence prior to
the southern supercontinent Gondwana breaking up.
The great diversification of marsupials in Australia and the absence of other mammals reflect Australia’s long isolation.
Australia has an abundance of endemic species—species found nowhere else—which is typical of islands whose isolation
by expanses of water prevents species to migrate. Over time, these species diverge evolutionarily into new species that
look very different from their ancestors that may exist on the mainland. The marsupials of Australia, the finches on the
Galápagos, and many species on the Hawaiian Islands are all unique to their one point of origin, yet they display distant
relationships to ancestral species on mainlands.
Molecular Biology
Like anatomical structures, the structures of the molecules of life reflect descent with modification. Evidence of a common
ancestor for all of life is reflected in the universality of DNA as the genetic material and in the near universality of
the genetic code and the machinery of DNA replication and expression. Fundamental divisions in life between the three
domains are reflected in major structural differences in otherwise conservative structures such as the components of
ribosomes and the structures of membranes. In general, the relatedness of groups of organisms is reflected in the similarity
of their DNA sequences—exactly the pattern that would be expected from descent and diversification from a common
ancestor.
DNA sequences have also shed light on some of the mechanisms of evolution. For example, it is clear that the evolution of
new functions for proteins commonly occurs after gene duplication events that allow the free modification of one copy by
mutation, selection, or drift (changes in a population’s gene pool resulting from chance), while the second copy continues
to produce a functional protein.

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Chapter 18 | Evolution and the Origin of Species 475

Misconceptions of Evolution
Although the theory of evolution generated some controversy when it was first proposed, it was almost universally accepted
by biologists, particularly younger biologists, within 20 years after publication of On the Origin of Species. Nevertheless,
the theory of evolution is a difficult concept and misconceptions about how it works abound.

This site (http://openstaxcollege.org/l/misconceptions) addresses some of the main misconceptions associated with the
theory of evolution.

Evolution Is Just a Theory

Critics of the theory of evolution dismiss its importance by purposefully confounding the everyday usage of the word
“theory” with the way scientists use the word. In science, a “theory” is understood to be a body of thoroughly tested and
verified explanations for a set of observations of the natural world. Scientists have a theory of the atom, a theory of gravity,
and the theory of relativity, each of which describes understood facts about the world. In the same way, the theory of
evolution describes facts about the living world. As such, a theory in science has survived significant efforts to discredit
it by scientists. In contrast, a “theory” in common vernacular is a word meaning a guess or suggested explanation; this
meaning is more akin to the scientific concept of “hypothesis.” When critics of evolution say evolution is “just a theory,”
they are implying that there is little evidence supporting it and that it is still in the process of being rigorously tested. This
is a mischaracterization.
Individuals Evolve
Evolution is the change in genetic composition of a population over time, specifically over generations, resulting from
differential reproduction of individuals with certain alleles. Individuals do change over their lifetime, obviously, but this is
called development and involves changes programmed by the set of genes the individual acquired at birth in coordination
with the individual’s environment. When thinking about the evolution of a characteristic, it is probably best to think about
the change of the average value of the characteristic in the population over time. For example, when natural selection leads
to bill-size change in medium-ground finches in the Galápagos, this does not mean that individual bills on the finches are
changing. If one measures the average bill size among all individuals in the population at one time and then measures the
average bill size in the population several years later, this average value will be different as a result of evolution. Although
some individuals may survive from the first time to the second, they will still have the same bill size; however, there will be
many new individuals that contribute to the shift in average bill size.
Evolution Explains the Origin of Life
It is a common misunderstanding that evolution includes an explanation of life’s origins. Conversely, some of the theory’s
critics believe that it cannot explain the origin of life. The theory does not try to explain the origin of life. The theory of
evolution explains how populations change over time and how life diversifies the origin of species. It does not shed light on
the beginnings of life including the origins of the first cells, which is how life is defined. The mechanisms of the origin of
life on Earth are a particularly difficult problem because it occurred a very long time ago, and presumably it just occurred
once. Importantly, biologists believe that the presence of life on Earth precludes the possibility that the events that led to
life on Earth can be repeated because the intermediate stages would immediately become food for existing living things.
However, once a mechanism of inheritance was in place in the form of a molecule like DNA either within a cell or pre-cell,
these entities would be subject to the principle of natural selection. More effective reproducers would increase in frequency
at the expense of inefficient reproducers. So while evolution does not explain the origin of life, it may have something to
say about some of the processes operating once pre-living entities acquired certain properties.
Organisms Evolve on Purpose
Statements such as “organisms evolve in response to a change in an environment” are quite common, but such statements
can lead to two types of misunderstandings. First, the statement must not be understood to mean that individual organisms
evolve. The statement is shorthand for “a population evolves in response to a changing environment.” However, a
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second misunderstanding may arise by interpreting the statement to mean that the evolution is somehow intentional.
A changed environment results in some individuals in the population, those with particular phenotypes, benefiting and
therefore producing proportionately more offspring than other phenotypes. This results in change in the population if the
characteristics are genetically determined.
It is also important to understand that the variation that natural selection works on is already in a population and does not
arise in response to an environmental change. For example, applying antibiotics to a population of bacteria will, over time,
select a population of bacteria that are resistant to antibiotics. The resistance, which is caused by a gene, did not arise by
mutation because of the application of the antibiotic. The gene for resistance was already present in the gene pool of the
bacteria, likely at a low frequency. The antibiotic, which kills the bacterial cells without the resistance gene, strongly selects
individuals that are resistant, since these would be the only ones that survived and divided. Experiments have demonstrated
that mutations for antibiotic resistance do not arise as a result of antibiotic.
In a larger sense, evolution is not goal directed. Species do not become “better” over time; they simply track their changing
environment with adaptations that maximize their reproduction in a particular environment at a particular time. Evolution
has no goal of making faster, bigger, more complex, or even smarter species, despite the commonness of this kind of
language in popular discourse. What characteristics evolve in a species are a function of the variation present and the
environment, both of which are constantly changing in a non-directional way. What trait is fit in one environment at one
time may well be fatal at some point in the future. This holds equally well for a species of insect as it does the human
species.

18.2 | Formation of New Species

By the end of this section, you will be able to:
• Define species and describe how species are identified as different
• Describe genetic variables that lead to speciation
• Identify prezygotic and postzygotic reproductive barriers
• Explain allopatric and sympatric speciation
• Describe adaptive radiation

Although all life on earth shares various genetic similarities, only certain organisms combine genetic information by sexual
reproduction and have offspring that can then successfully reproduce. Scientists call such organisms members of the same
biological species.

Species and the Ability to Reproduce

A species is a group of individual organisms that interbreed and produce fertile, viable offspring. According to this
definition, one species is distinguished from another when, in nature, it is not possible for matings between individuals from
each species to produce fertile offspring.
Members of the same species share both external and internal characteristics, which develop from their DNA. The closer
relationship two organisms share, the more DNA they have in common, just like people and their families. People’s DNA
is likely to be more like their father or mother’s DNA than their cousin or grandparent’s DNA. Organisms of the same
species have the highest level of DNA alignment and therefore share characteristics and behaviors that lead to successful
reproduction.
Species’ appearance can be misleading in suggesting an ability or inability to mate. For example, even though domestic
dogs (Canis lupus familiaris) display phenotypic differences, such as size, build, and coat, most dogs can interbreed and
produce viable puppies that can mature and sexually reproduce (Figure 18.9).

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