CHAPTER 14

The Origin of Species

Chapter Objectives
Opening Essay
Explain how speciation may be occurring in the Vogelkop bowerbird.

Defining Species
14.1 Distinguish between microevolution and speciation.
14.2 Compare the definitions, advantages, and disadvantages of the different species
concepts.
14.3 Describe five types of prezygotic barriers and three types of postzygotic
barriers that prevent populations of closely related species from interbreeding.

Mechanisms of Speciation
14.4 Explain how geologic processes can fragment populations and lead to speciation.
14.5 Explain how reproductive barriers might evolve in isolated populations of
organisms. Refer to studies of laboratory-raised fruit flies and monkey flowers.
14.6 Explain how sympatric speciation can occur, noting examples in plants and
animals.
14.7 Explain why polyploidy is important to modern agriculture. Explain how
modern wheat evolved.
14.8 Describe the circumstances that led to the adaptive radiation of the Galápagos
finches.
14.9 Explain how coloration and properties of light transmission in water has led to
speciation in Lake Victoria.
14.10 Explain how hybrid zones affect speciation. Describe examples of
reinforcement and fusion in hybrid zones.
14.11 Compare the gradual model and the punctuated equilibrium model of evolution.
Explain how each model applies to the fossil record.

Lecture Outline
I. Introduction
1. Bowerbirds, native to New Guinea and Australia, are named for the structure called a
bower that the male weaves from twigs and grasses to attract females.
2. After building his bower, the male collects objects such as fruits, seeds, insect parts,
rocks, flowers, and leaves and arranges them artfully by color and type.
3. Females
a. tour the bowers of local males, inspecting each bower carefully while its owner
courts her with a song and dance, and

Copyright © 2015 Pearson Education, Inc. 187

 b. may visit promising candidates multiple times before finally mating with one.
4. Not all Vogelkop bowerbirds construct displays.
5. The males of another population build a simpler structure consisting of sticks loosely
woven around a central sapling.
6. Researchers hypothesize that this divergence in display preferences has started the two
populations on separate evolutionary paths, with each path leading to a new species, or
speciation.
7. An ancestral cormorant species is thought to have flown from the Americas to the
Galápagos Islands more than 3 million years ago.
8. Terrestrial mammals could not make the trip over the wide distance, and no predatory
mammals naturally occur on these islands today.
9. Without predators, the environment of these cormorants favored birds with smaller
wings, perhaps channeling resources to the production of offspring.
II. Defining Species
A. 14.1 The origin of species is the source of biological diversity
1. Darwin was eager to explore landforms newly emerged from the sea when he came to
the Galápagos Islands.
2. He noted that these volcanic islands, despite their geologic youth, were teeming with
plants and animals found nowhere else in the world.
3. He realized that these species, like the islands, were relatively new.
4. Microevolution is the change in the gene pool of a population from one generation to
the next.
5. Speciation is the process by which one species splits into two or more species.
6. Each time speciation occurs, the diversity of life increases.
7. Over the course of 3.5 million years,
a. an ancestral species first gave rise to two or more different species,
b. which then branched to new lineages,
c. which branched again,
d. until we arrive at the millions of species that live, or once lived, on Earth.
B. 14.2 There are several ways to define a species
1. The word species is from the Latin for “kind” or “appearance.”
2. Although the basic idea of species as distinct life-forms seems intuitive, devising a
more formal definition is not easy and raises questions.
3. In many cases, the differences between two species are obvious. In other cases, the
differences between two species are not so obvious.
4. How similar are members of the same species?
a. Whereas the individuals of many species exhibit fairly limited variation in physical
appearance, certain other species—our own, for example—seem extremely varied.
5. The biological species concept defines a species as a group of populations, whose
members have the potential to interbreed in nature and produce fertile offspring
(offspring that themselves can reproduce).
6. Thus, members of a biological species are united by being reproductively compatible,
at least potentially.
7. Reproductive isolation
a. prevents genetic exchange (gene flow) and
b. maintains a boundary between species.

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 8. But there are some pairs of clearly distinct species that do occasionally interbreed.
a. The resulting offspring are called hybrids.
b. An example is the grizzly bear (Ursus arctos) and the polar bear (Ursus maritimus),
whose offspring have been called “grolar bears.”
9. There are other instances in which applying the biological species concept can be
problematic.
a. There is no way to determine whether organisms that are now known only through
fossils were once able to interbreed.
b. Reproductive isolation does not apply to prokaryotes or other organisms that
reproduce only sexually.
c. Therefore, alternative species concepts can be useful.
10. The morphological species concept
a. classifies organisms based on observable physical traits and
b. can be applied to asexual organisms and fossils.
11. However, there is some subjectivity in deciding which traits to use.
12. The ecological species concept
a. defines a species by its ecological role or niche and
b. focuses on unique adaptations to particular roles in a biological community.
13. For example, two species may be similar in appearance but distinguishable
based on what they eat or the depth of water in which they are usually found.
14. The phylogenetic species concept defines a species as the smallest group of
individuals that share a common ancestor and thus form one branch of the tree of life.
15. Biologists trace the phylogenetic history of a species by comparing its
a. morphology,
b. DNA sequences, or
c. biochemical pathways.
16. However, defining the amount of difference required to establish separate species
remains a challenge.
C. 14.3 VISUALIZING THE CONCEPT: Reproductive barriers keep species separate
1. Reproductive barriers
a. serve to isolate the gene pools of species and
b. prevent interbreeding.
2. Depending on whether they function before or after zygotes form, reproductive
barriers are categorized as
a. prezygotic or
b. postzygotic.
3. Five types of prezygotic barriers prevent mating or fertilization between species.
a. In habitat isolation, there is a lack of opportunity for mates to encounter each other.
b. In temporal isolation, there is breeding at different times or seasons.
c. In behavioral isolation, there is failure to send or receive appropriate signals.
d. In mechanical isolation, there is a physical incompatibility of reproductive parts.
e. In gametic isolation, there is molecular incompatibility of eggs and sperm or pollen
and stigma.
4. Three types of postzygotic barriers operate after hybrid zygotes have formed.
a. In reduced hybrid viability, interaction of parental genes impairs the hybrid’s
development or survival.
b. In reduced hybrid fertility, hybrids are vigorous but cannot produce viable
offspring.

Copyright © 2015 Pearson Education, Inc. CHAPTER 14 The Origin of Species 189
 c. In hybrid breakdown, hybrids are viable and fertile, but their offspring are feeble or
sterile.
III. Mechanisms of Speciation
A. 14.4 In allopatric speciation, geographic isolation leads to speciation
1. A key event in the origin of a new species is the separation of a population from other
populations of the same species.
a. With its gene pool isolated, the splinter population can follow its own evolutionary
course.
b. Changes in allele frequencies caused by natural selection, genetic drift, and
mutation will now be diluted by alleles entering from other populations (gene flow).
2. In allopatric speciation, the initial block to gene flow may come from a geographic
barrier that isolates a population.
3. Several geologic processes can isolate populations.
a. A mountain range may emerge and gradually split a population of organisms that
can inhabit only lowlands.
b. A large lake may subside until there are several smaller lakes, isolating certain fish
populations.
c. Continents themselves can split and move apart.
d. Allopatric speciation can also occur when individuals colonize a remote area and
become geographically isolated from the parent population.
4. How large must a geographic barrier be to keep allopatric populations apart?
a. The answer depends on the ability of the organisms to move.
b. Birds, mountain lions, and coyotes can easily cross mountain ranges.
c. In contrast, small rodents may find a canyon or a wide river a formidable barrier.
The Grand Canyon and Colorado River separate two species of antelope squirrels.
5. Thirty species of snapping shrimp in the genus Alpheus live off the Isthmus of
Panama, the land bridge that connects South and North America.
a. Morphological and genetic data group these shrimp into 15 pairs of species, with
the members of each pair being each other’s closest relative.
b. In each case, one member of the pairs lives on the Atlantic side of the isthmus,
whereas the other lives on the Pacific side.
c. This strongly suggests that geographic separation of the ancestral species of these
snapping shrimp led to allopatric speciation.
B. 14.5 Reproductive barriers can evolve as populations diverge
1. How do reproductive barriers arise?
2. The environment of an isolated population my include
a. different food sources,
b. different types of pollinators, and
c. different predators.
3. As a result of natural selection acting on preexisting variations (or as a result of
genetic drift or mutation), a population’s traits may change in ways that also establish
reproductive barriers.
4. Researchers have successfully documented the evolution of reproductive isolation
with laboratory experiments.
5. These studies have included
a. laboratory studies of fruit flies and
b. field studies of monkey flowers and their pollinators.

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 C. 14.6 Sympatric speciation takes place without geographic isolation
1. Sympatric speciation occurs when a new species arises within the same geographic
area as a parent species.
a. How can reproductive isolation develop when members of sympatric populations
remain in contact with each other?
2. Gene flow between populations may be reduced by
a. polyploidy,
b. habitat differentiation, or
c. sexual selection.
3. Many plant species have originated from sympatric speciation that occurs when
accidents during cell division result in extra sets of chromosomes.
4. New species formed in this way are polyploid, in that their cells have more than two
complete sets of chromosomes.
5. Sympatric speciation can result from polyploidy
a. within a species (by self-fertilization) or
b. between two species (by hybridization).
D. 14.7 EVOLUTION CONNECTION: The origin of most plant species can be traced to
polyploid speciation
1. Plant biologists estimate that 80% of all living plant species are descendants of
ancestors that formed by polyploid speciation.
2. Hybridization between two species accounts for most of these species, perhaps
because of the adaptive advantage of the diverse genes a hybrid inherits from different
parental species.
3. Polyploid plants include
a. cotton,
b. oats,
c. potatoes,
d. bananas,
e. peanuts,
f. barley,
g. plums,
h. apples,
i. sugarcane,
j. coffee, and
k. wheat.
4. Wheat
a. has been domesticated for at least 10,000 years and
b. is the most widely cultivated plant in the world.
5. Bread wheat, Triticum aestivum, is
a. a polyploid with 42 chromosomes and
b. the result of hybridization and polyploidy.
E. 14.8 Isolated islands are often showcases of speciation
1. Isolated island chains are often inhabited by unique collections of species.
2. Islands that have physically diverse habitats and that are far enough apart to permit
populations to evolve in isolation but close enough to allow occasional dispersions to
occur are often the sites of multiple speciation events.
3. The evolution of many diverse species from a common ancestor is known as adaptive
radiation.

Copyright © 2015 Pearson Education, Inc. CHAPTER 14 The Origin of Species 191
 4. The Galápagos Archipelago
a. is located about 900 km (560 miles) west of Ecuador,
b. is one of the world’s great showcases of adaptive radiation,
c. was formed naked from underwater volcanoes from 5 million to 1 million years
ago,
d. was colonized gradually from other islands and the South America mainland, and
e. has many species of plants and animals found nowhere else in the world.
5. The Galápagos Islands currently have 14 species of closely related finches, called
Darwin’s finches, because Darwin collected them during his around-the-world voyage
on the Beagle.
6. These birds
a. share many finchlike traits,
b. differ in their feeding habits and their beaks, specialized for what they eat, and
c. arose through adaptive radiation.
F. 14.9 SCIENTIFIC DISCOVERY: A long-term field study documents evolution in
Darwin’s finches
1. We can see speciation occurring.
2. The species living today represent a snapshot, a brief instant in this vast span of time.
3. The environment continues to change, sometimes rapidly due to human impact, and
natural selection continues to act on affected populations.
4. Researchers have documented at least two dozen cases in which populations are
diverging as they exploit different food resources or breed in different habitats.
5. Sexual selection is a form of natural selection in which individuals with certain traits
are more likely to obtain mates.
6. In addition to the bowerbirds already discussed, biologists have identified several
other animal populations that are diverging as a result of differences in how males
attract females or how females choose mates.
7. Biologists can also test hypotheses about the process of speciation by studying species
that arose recently.
8. Cichlids are a family of fishes that live in tropical lakes and rivers.
a. The come in all colors of the rainbow.
b. They are renowned for the spectacular adaptive radiations that stocked the large
lakes of East Africa with more than a thousand species of cichlids in less than
100,000 years.
c. In the largest of these lakes, Lake Victoria, roughly 500 species evolved in about
15,000 years.
9. In Lake Victoria, there are pairs of closely related cichlid species that differ in color
but nothing else.
a. Breeding males of Pundamilia nyererei have a bright red back and dorsal fin.
b. Breeding males of Pundamilia pundamilia are metallic blue-gray.
10. Pundamilia females prefer brightly colored males.
11. Mate-choice experiments performed in the laboratory showed that
a. P. nyererei females prefer red males over blue males,
b. P. pundamilia females prefer blue males over red males,
c. the vision of P. nyererei females is more sensitive to red light than blue light, and
d. the vision of P. pundamilia females is more sensitive to blue light than red light.
12. Researchers also demonstrated that this color sensitivity is heritable.

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 13. As light travels through water, suspended particles selectively absorb and scatter the
shorter (blue) wavelengths, so light becomes increasingly red with increasing depth.
14. Thus, in deeper waters, P. nyererei males are pleasingly apparent to females with
red-sensitive vision but virtually invisible to P. pundamilia females.
15. When biologists sampled cichlid populations in Lake Victoria, they found that
a. P. neyererei breeds in deep water, whereas
b. P. pundamilia inhabits shallower habitats where the blue males shine brightly.
16. As a consequence of their mating behavior, the two species encounter different
environments that may result in further divergence.
G. 14.10 Hybrid zones provide opportunities to study reproductive isolation
1. What happens when separated populations of closely related species come back into
contact with each other?
2. Biologists try to answer such questions by studying hybrid zones, regions in which
members of different species meet and mate to produce at least some hybrid offspring.
3. Figure 14.10A illustrates the formation of a hybrid zone, starting with the ancestral
species.
4. Reinforcement
a. When hybrid offspring are less fit than members of both parent species, we might
expect
i. natural selection to strengthen, or reinforce, reproductive barriers, thus reducing
the formation of unfit hybrids, and
ii. that barriers between species should be stronger where the species overlap
(that is, where the species are sympatric).
b. The closely related collared flycatcher and pied flycatcher are an example of
reinforcement.
5. Fusion
a. What happens when the reproductive barriers between species are not strong and
the species come into contact in a hybrid zone?
b. So much gene flow may occur that the speciation process reverses, causing the two
hybridizing species to fuse into one.
c. Such a situation has been occurring among the cichlid species in Lake Victoria.
d. Pollution caused by development along the shores of Lake Victoria has turned the
water murky.
e. What happens when P. nyererei or P. pundamilia females can’t tell red males from
blue males?
i. The behavior barrier crumbles.
ii. Many viable hybrid offspring are produced by interbreeding.
iii. The once isolated gene pools of the parent species are combining, with two
species fusing into a single hybrid species.
H. 14.11 Speciation can occur rapidly or slowly
1. There are two models for the tempo of speciation.
a. The punctuated equilibria model draws on the fossil record, where species change
most as they arise from an ancestral species and then change relatively little for the
rest of their existence.
b. Other species appear to have evolved more gradually.
2. The time interval between speciation events varies from a few thousand years to tens
of millions of years.

Copyright © 2015 Pearson Education, Inc. CHAPTER 14 The Origin of Species 193
Chapter Guide to Teaching Resources

Defining Species (14.1–14.3)

Student Misconceptions and Concerns
• Students might not realize that evolutionary change includes both (a) linear events, in
which a species changes over time, and (b) branching events, which produce new species
and diversity. Some students simply expect that whenever new species evolve, they
replace their ancestors. (14.1)
• Students might have never considered how species are naturally kept separate and unique.
Instead, students may consider species as fixed entities, especially the species to which
they belong. To help ease students into the topic, consider pointing out that species do not
reflect an even spectrum of diversity. Instead, there are many groups of clearly related
organisms (owls, grasses, sharks, beetles, butterflies, trees, mushrooms, and bacteria, for
example). Ask students to consider why such groupings exist. Could such groupings
represent shared ancestry? (14.2–14.3)

Teaching Tips
• Challenge your students to explain why the field of paleontology has largely been
concerned with macroevolution. The broader perspective of evolutionary change studied
by paleontologists rarely permits an examination of change within a species. (14.1)
• Identify or have your students find several commonly recognized and related species of
plants or animals in your area and find out what reproductive barriers keep these species
from interbreeding. Local examples always help to bring a point home. (14.3)

Active Lecture Tips

• Before lecturing about species concepts, consider a short writing assignment. Have
students work individually or in small groups, without the benefit of books, to define a
species. (14.2)

Mechanisms of Speciation (14.4–14.11)

Student Misconceptions and Concerns
• Students must understand that species do not evolve because of need. Biological diversity
exists and the environment selects. Evolution is not deliberate; it is reactive. Species do not
deliberately change. There is no plan. As teachers, we must take care that our descriptions
of evolution accurately reflect its process. The use of the passive voice in descriptions of
evolution is one way of doing this. (14.4–14.11)
• Most of us are unable to comprehend the vast lengths of time considered by geologists.
Exercises and examples can increase this comprehension. Consider the number of seconds
in a year (60 × 60 × 24 × 365.25 = 31,557,600) or how much money you could spend each
day if you spent $1 million a year ($1,000,000/365 = $2,739.73/day). (14.4–14.11)
• Students also need to be reminded that 1 billion is 1,000 million. Many students (and some
politicians) easily confuse million and billion without realizing the scale of the error.
(14.4–14.11)

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 • The concept of rarity is likely to be misunderstood when applied to geologic time. Events
such as major floods, earthquakes, or asteroid impacts, which might be so rare as to occur
every 1,000 years, are actually common in geological terms. Students might not realize
that 1,000 such events would be expected to occur over a million years. (14.11)

Teaching Tips
• The isolation of a few individuals from a parent population may result from a catastrophic
weather or geological event. Ask your students to think back to news footage of torrential
rains, massive debris rocketing down a river, and the struggles of animals to haul
themselves onto these rafts. Better yet, show them a short news clip of such events.
Dramatic weather and geological events may be rare in our lifetimes but are frequent
enough to play a role in speciation. (14.4)
• When discussing Module 14.5, consider referring back to Figure 14.3A. Challenge
students to explain how each of the prezygotic barriers might impact the evolution of a
new species. (14.5)
• The Silvery Salamander, Ambystoma platineum, is a triploid, all-female species living in
parts of the U.S. Midwest. It is believed to have formed by the hybridization of two related
species thousands of years ago. It is an unusual example of sympatric speciation in
animals. A good starting point for learning more about this species is
www.inhs.illinois.edu/animals_plants/herps/species/am_platine.html. (14.6)
• The abundance of polyploid plants used for food, facilitates further study for student
assignments. Perhaps small groups or individuals can select a polyploid crop and describe
its evolutionary history and/or its current method of reproduction. (14.7)
• An analogy might be made between the specialized functions of finch beaks and the many
types of screwdrivers (or pliers) that exist today. Each type of screwdriver (Phillips,
flathead, hex, etc.) represents a specialization for a particular job or a generalist approach,
useful in a variety of applications. (14.8)
• Numerous examples of adaptive radiations exist in the Hawaiian Islands. Hawaiian
honeycreepers (birds), fruit flies, and species of the plant genera Cyrtandra and Geranium
are excellent examples for additional illustration. (14.8)
• The work of Peter and Rosemary Grant with Darwin’s finches helps to explain to students
that the concept of “better” in evolution is relative. As the environment changes, organisms
must respond or suffer the consequences. In these circumstances, organisms are reacting,
not improving. (14.9)
• Students might wish to debate whether two cichlid species that fuse into one were
previously separate species. If each species retained the natural ability to hybridize with
each other, and did so extensively as the environment changed, were they separate species?
Such difficult distinctions test our definitions and reveal some of the challenges of biology.
(14.10)

Active Lecture Tips

• Have your students work in pairs to think of analogous examples of punctuated
equilibrium in our culture. One such example is the switch from vinyl records to compact
discs, with the brief transitional form of cassette tapes (which students currently entering
college may barely remember). Between the years 1900 and 2000, there were both long
periods of stasis (vinyl records) and a relatively short period of transition to CDs and now

Copyright © 2015 Pearson Education, Inc. CHAPTER 14 The Origin of Species 195
 to digital music files (who knows how long they will last?). Similarly, black-and-white TV
to color or high-definition and soon 3-D represent similar punctuated changes. Debating
the validity of analogies can itself be instructive as students articulate the biological
principles and compare them to the analogies. (14.11)
• See the Activity, Video Review: YouTube – Bang Goes the Theory: Evolution Made
Simply, at the Instructor Exchange in the Mastering Biology instructor resource area or at
the following YouTube address: www.youtube.com/watch?v=YYLKt5J-
KKY&noredirect=1. (14.4–14.11)

Key Terms
adaptive radiation phylogenetic species concept
allopatric speciation polyploid
biological species postzygotic barriers
concept prezygotic barriers
ecological species punctuated equilibria
concept reproductive isolation
hybrids speciation
hybrid zone species
morphological species sympatric speciation
concept

Word Roots
allo- = other; -patri = father (allopatric speciation: the formation of new species in populations
that are geographically isolated from one another)
morpho = form (morphological species concept: classification based mainly on physical traits
such as shape, size, and other features of morphology)
poly- = many (polyploid: cells with more than two complete sets of chromosomes)
post- = after (postzygotic barrier: a reproductive barrier that prevents hybrid zygotes produced by
two different species from developing into viable, fertile adults)
pre- = before (prezygotic barrier: a reproductive barrier that impedes mating between species or
hinders fertilization if mating between two species is attempted)
sym- = together; -patri = father (sympatric speciation: the formation of new species in populations
that live in the same geographic area)

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