Iguanas as Examples of Evolutionary Change

guanas are a type of lizard known for a very long tail, which can be up to
three times their body length. The green iguana is commo'i'rthroughout the
South American continent. Its claws enable it to climb trees, where it feeds on
succulent leaves and soft fruits along riverbanks. It is an excellent swimmer
and uses rivers to travel to new feeding areas.
Green iguanas do not inhabit islands to the west and northeast of South
America. Instead, the Galapagos Islands to the west have one type of land
iguana and one type of marine iguana. Marine iguanas, black in color, are
unique to the Galapagos Islands-they occur no place else on Earth. These
air-breathing reptiles can dive to a depth exceeding 10 meters (33 feet) and
remain submerged for more than 30 minutes. They use their claws to cling to
the bottom rocks while feeding on algae.
Hispaniola, an island to the northeast of South America, is inhabited by
the rhinoceros iguana, which has three horny bumps on its snout and is dark
brown to black in color. This iguana lives mainly in dry forests with rocky,
lmestone habitats and feeds on a wide variety of plants.
The theory of evolution explains the occurrence of the unique iguanas
on the Galapagos Islands and Hispaniola. It is hypothesized and supported
by various data that ancestral iguanas swam or hitchhiked on floating drift-
wood from South America to the Galapagos to the west and to Hispaniola to
the northeast. After arrival, these iguana populations were cut off from other
iguana populations, and this allowed them to evolve into these new and dif-
ferent species. The origin of species and how they are classified and studied
are the topics of this chapter.
Evolution on
a Large Scale
OUTLINE
16.1 Speciation and Macroevolution 266
16.2 The Fossil Record 272
16.3 Systematics 277
BEFORE YOU BEGIN
Before beginning this chapter, take a few moments to
review the following discussions.
Section 14.2 What roles do the fossil records and the
study of comparative anatomy have in understanding
evolutionary change?
Section 15.1 How does natural selection act as the
mechanism of evolutionary change?
Section 15.2 What is microevolution?
265
266 PARTTHREE Evolution
Figure 16.1 Dinosaurs.
Artwork of a Mesozoic landscape, Cretaceous period,
including saltasaurus, brachiosaurus, mononykus,
protoceratops, nyctosaurus, styracosaurus, tarbosaurus,
maiasaura, stegoceras, evoplocephalus, deltatheridium, and
ichthyornis (145 to 65 MYA). All of these species are now extinct.
16.1 Speciation and Macroevolution
Learning Outcomes
Upon completion of this section, you should be able to
1. Explain how scientists define a biological species and describe the
limitation of this definition.
2. List five types of prezygotic isolating mechanisms and explain how
they prevent members of two different species from reproducing.
3. List three types of postzygotic isolating mechanisms and explain
how they prevent members of two different species from
reproducing.
4. Contrast allopatric speciation with sympatric speciation and explain
how each method may result in the creation of a new species.
5. Explain how new species may arise by adaptive radiation.
Chapter 15 considered evolution on a small scale-that is, microevolution,
small changes over a short period of time. In this chapter, we turn our attention
to evolution on a large scale-that is, macroevolution, large changes over a
very long period of time. The history of life on Earth is a part of macroevolu-
tion. Macroevolution requires speciation, the splitting of one species into two
or more new species. Speciation involves the gene pool changes that we studied
in Chapter 15.
Species originate, evolve adaptations to their environments, and then may
become extinct (Fig. 16.1). Without the origin and extinction of species, life on
Earth would not have the ever-changing history that we find in the fossil record.
)
Defining Species
Before we consider the origin of species, we first need to define a species.
Appearance is not always a good criterion. The members of different spe-
cies can look quite similar, while the members of a si ngle species can be
diverse in appearance. Although many definitions have been proposed, the
.ion,
tion
era
olu-
two
lied
ay
on
)rd.
ies.
e-
be

hio/ogical species concept offers a testable way to define a species that does not
depend on appearance: The members of a species interbreed and have a shared
gene pool, and each species is reproductively isolated from every other species.
For example, the flycatchers in Figure 16.2 are members of separate species
because they do not interbreed in nature.
According to the biological species concept, gene flow occurs between
the populations of a species, but not between populations of different species.
The red maple and the sugar maple are found over a wide geographic range
m the eastern half of the United States, and each species is made up of many
populations. However, the members of each species' population rarely hybridize
m nature. Therefore, these two types of plants are separate species. In contrast,
the human species has many populations, which certainly differ in physical
appearance (Fig. 16.3). We know, however, that all humans belong to one spe-
cies because the members of these populations can produce fertile offspring.
The biological species concept is useful, as we shall
see, but even so, it has its limitations. For example,
it applies only to sexually reproducing organ-
Isms and cannot apply to asexually reproducing
organisms. Then, too, sexually reproducing
organisms are not always as reproductively
1solated as we would expect. Some North
orioles live in the western half of
the continent, some in the eastern half, yet even
the two most genetically distant oriole species, as
recognized by analyzing their mitochondrial DNA,
will hybridize where they meet in the middle of the
continent.
There are other definitions of
1pecies aside from the biological
definition. Later in this chapter, we
will define species as a category
of classification below the rank of
genus. Species in the same genus
1hare a recent common ancestor.
Empidonax trailli
CHAPTER 16 Evolution on a Large Scale
Empidonax virescens
Least flycatcher,
Empidonax minimus
Figure 16.2 Three species of flycatchers.
Although these flycatcher species are nearly identical in appearance,
we know they are separate species because they are reproductively
isolated-the members of each species reproduce only with one
another. Each species has a characteristic song and its own habitat
during the mating season as well.
Figure 16.3 Human populations.
a. b.
The Maassai of East Africa (a) and the Kuna Indians from the San Bias Islands of Panama
(b) are both members of the species Homo sapiens because the Maassai and Kuna can
reproduce and produce fertile offspring.
268 PART THREE Evolution
Connections and Misconceptions
How can we determine if an organism that does not
reproduce sexually is a distinct species?
Many organisms either do not or
very rarely reproduce sexually.
For example, there are species of
mosses that reproduce sexually only
every 200 to 300 years! To determine
if two populations of asexual organ-
isms are distinct species, scientists
rely on DNA analysis, morphological
studies, and a close examination of the organisms' ecology
to determine whether the two populations would reproduce
naturally. Often, scientists have to revisit the classification of
a species as research unveils new information.
A common ancestor is a single ancestor for two or more different groups. For
example, your father's mother is the common ancestor for you, your siblings.
and your paternal cousins. Similarly, there is a common ancestor for all species
of roses.
Reproductive Barriers
As mentioned, for two species to be separate, they must be reproductively
isolated-that is, gene flow must not occur between them. Reproductive barri
ers are isolating mechanisms that prevent successful reproduction (Fig. 16.4).
In evolution, reproduction is successful only when it produces fertile offspring.
Prezygotic (before the formation of a zygote) isolating mechanisms
prevent reproductive attempts and make it unlikely that fertilization will be sue
cessful if mating is attempted. Habitat isolation, temporal isolation, behavioral
isolation, mechanical isolation, and gamete isolation make it highly unlikely that
particular genotypes will contribute to the gene pool of a population.
Habitat isolation When two species occupy different habitats, even within
the same geographic range, they are less likely to meet and attempt
to reproduce. This is one of the reasons that the flycatchers in Figure
16.2 do not mate and the red maple and sugar maple do not exchange
pollen. In tropical rain forests, many animal species are restricted to a
particular level of the forest canopy; in this way, they are isolated from
similar species.
Temporal isolation Two species can live in the same locale, but if they
reproduce at different times of year, they do not attempt to mate. For
example, Reticulitermes hageni and R. virginicus are two species of
termites. The former has mating flights in March through May, whereas
the latter mates in the fall and winter months .
. ,.
Prezygotlc Isolating mechanisms Postzygotic isolating mechanisms
Premating
Habitat isolation
Species at same locale occupy
different habitats.
Temporal isolation
Species reproduce at different
seasons or different times
of day.
Behavioral isolation
In animal species, courtship
behavior differs, or individuals
respond to different songs, calls,
pheromones, or other signals.
Figure 16.4 Reproductive barriers.
Mating
Mechanical isolation
Genital ia between
species are unsuitable
for one another.
Gamete isolation
Sperm cannot reach or
fertilize egg.
Fertilization
Zygote mortality
Fertilization occurs, but
zygote does not survive.
Hybrid sterility
Hybrid survives but is sterile
and cannot reproduce.
F
2
fitness
Hybrid is fertile, but F
2
hybrid
has reduced fitness.
Prezygotic isolating mechanisms prevent mating attempts or a successful outcome, should mating take place-for example, between two species of orioles. No zygote is
ever formed if these mechanisms are successful. Postzygotic isolating mechanisms prevent offspring from reproducing- that is, if a hybrid oriole should result, it would be
unable to breed successfully.
Behavioral isolation Many anima! species have courtship patterns that allow
males and females to recognize one another (Fig. 16.5). Female fireflies
recognize males of their species by the pattern of the males' flashings;
similarly, female crickets recognize males of their species by the males'
chirping. Many males recognize females of their species by sensing
chemical signals, called pheromones. For example, female gypsy moths
secrete chemicals from abdominal glands. These
chemicals are detected downwind by receptors on
the antennae of males.
Video
Flirting Flies
Mechanical isolation When animal genitalia or plant floral structures are
incompatible, reproduction cannot occur. Inaccessibility of pollen to
certain pollinators can prevent cross-fertilization in plants, and the sexes
of many insect species have genitalia that do not match, or other
characteristics that make mating impossible. For example, male
dragonflies have claspers that are suitable for holding only the females
of their own species.
Gamete isolation Even if the gametes of two different species meet, they may
not fuse to become a zygote. In animals, the sperm of one species may
not be able to survive in the reproductive tract of another species, or
the egg may have receptors only for sperm of its species. Also, in each
type of flower, only certain pollen grains can germinate, so that sperm
successfully reach the egg.
Postzygotic (after the formation of a zygote) isolating mechanisms prevent
hybrid offspring (reproductive product of two different species) from develop-
ing or breeding, even if reproduction attempts have been successful.
Zygote mortality The hybrid zygote may not be viable, so it dies. A zygote with
two different chromosome sets may fail to go through mitosis properly,
or the developing embryo may receive incompatible instructions from
the maternal and paternal genes, so that it cannot continue to exist.
Hybrid sterility The hybrid zygote may develop into a sterile adult. As is well
known, a cross between a horse and a donkey produces a mule, which is
usually sterile-it cannot reproduce (Fig. 16.6). Sterility of hybrids
generally results from complications in meiosis, which lead to an inability
to produce viable gametes. A cross between a cabbage and a radish
produces offspring that cannot form gametes, even though the diploid
number is 18, an even number, most likely because the cabbage
chromosomes and the radish chromosomes cannot align during meiosis.
F2funess If hybrids can reproduce, their offspring are unable to reproduce. In
some cases, mules are fertile, but their offspring (the F
2
generation) are
not fertile.
Models of Speciation
The introduction to this chapter uggests that iguanas of South America may be
the common ancestor for both the marine iguana on the Galapagos Islands (to the
west of South America) and the rhinoceros iguana on Hispaniola (the Caribbean
island containing the countries of Haiti and the Dominican Republic). If so, how
could it have happened? Green iguanas are strong swimmers, so by chance a few
could have migrated to these islands, where they formed populations separate
from each other and from the parent population in South America. Each popula-
tion continued on its own evolutionary path as new mutations, genetic drift, and
natural selection occurred. Eventually, reproductive isolation developed, and there
CHAPTER 16 Evolution on a Large Scale
Figure 16.5 Prezygotic isolating mechanism.
An elaborate courtship display allows the blue-footed boobies of thj
Galapagos Islands to select a mate. The male lifts his feet in a ritualiz
manner that shows off their bright blue color.
d'donkey
Figure 16.6 Postzygotic isolating mechanism.
Mules are horse-donkey hybrids. Mules are infertile due to a differenc
the chromosomes inherited from their parents.
270 PART THREE Evolution
were three species of iguanas. A speciation model based on geographic
isolation of populations is called allopatric speciation because al/o
means different and patria loosely means homeland.
Ensatina eschscholtzii
Figure 16.7 features an example of allopatric speciation thai
has been extensively studied in California. Members of an ancestral
population of Ensatina salamanders existing in the Pacific Northwe11
oregonensis
Western migration
southward establishes
these populations.
Ensatina eschscholtzii
xanthoptica
Eastern migration
southward establishes
these populations.
migrated southward, establishing a range of populations. Each population
was exposed to its own selective pressures along the coastal mountains and along
the Sierra Nevada Mountains. Due to the barrier created by the Central Valley of
California, limited gene flow occurred between the eastern populations and the
western populations. Genetic differences increased from north to south, resulting
in two distinct forms of Ensatina salamanders in Southern California that differ
dramatically in color and interbreed only rarely.
With sympatric speciation, a population develops into
two or more reproductively isolated groups without prior
geographic isolation. One of the best examples to eas
ily illustrate this type of speciation is found among
plants, where it can occur by means of polyploid':
an increase in the number of sets of chromosome'
Ensatina eschscholtzii
to 3n or above. The presence of sex chromosome1
makes it difficult for polyploidy speciation 10
occur in animals. In plants, hybridization between
two species can be followed by a doubling of the
chromosome number. Such polyploid planls
are reproductively isolated by a postzygotic
mechanism; they can reproduce successfully on!)
croceater
with other similar polyploids, and backcrosses
with their parents are sterile. Therefore, three species
instead of two species result. Figure 16.8 shows thai
the parents 9f our present-day bread wheat had 28
and 14, chromosomes, respectively. The
hybrid with 21 chromosomes is ster-
Ensatina eschscholtzii
eschschoftzii
In the end, two reproductively
isolated populations cannot
mate in Southern California.
Ensatina eschscholtzii
klauberi
ile, but polyploid bread wheat wilh
42 chromosomes is fertile because
the chromosomes can align during meiosis.
Adaptive Radiation
Figure 16.7 Allopatric speciation.
In this example of allopatric speciation, the Central Valley of California is
separating a range of populations descended from the same northern
ancestral species. The limited contact between the populations on the
west and those on the east allow genetic changes to build up to such an
extent that members of the two southern populations rarely reproduce
with each other and are designated as subspecies.
A clear example of speciation through adaptive radiation is pro-
vided by the finches on the Galapagos Islands, which are often called Darwin's
Figure 16.8 Sympatric speciation.
In this example of sympatric speciation, two populations of wild wheat hybridized
many years ago. The hybrid is sterile, but chromosome doubling allowed some plants to
reproduce. These plants became today's bread wheat.
X
Wild wheat
2n= 28
Triticum
turgidum
__..... Sterile hybrid
2n= 21
Wild wheat
2n= 14
Triticum
taushii
Doubling of
chromosome
number
Bread wheal
2n=42
Triticum
aestivum
finches because Darwin first realized their significance as an example of how
evolution works. During adaptive radiation, many new species evolve from a
single ancestral species. The many species of finches that live on the Galapagos
Islands are hypothesized to be descendants of a single type of ancestral finch from
the mainland (Fig. 16.9). The population on the various islands were subjected
to the founder effect involving genetic drift, genetic mutations, and the process of
natural selection. Because of natural selection, each population became adapted
to a particular habitat on its island. In time, the various populations became so
genotypically different that now, when by chance they reside on the same island,
they do not interbreed and are therefore separate species. There is evidence that
the finches use beak shape to recognize members of the same Video
species during courtship. Rejection of suitors with the wrong Finches
Adaptive
type of beak is a behavioral prezygotic isolating mechanism. Radiation
Similarly, inhabiting the Hawaiian Islands is a wide variety of honeycreep-
ers, all descended from a common goldfinch- like ancestor that arrived from Asia
or North America about 5 MYA. Today, honeycreepers have a range of beak sizes
and hapes (see Fig. I .11) for feeding on various food sources, including seeds,
fruits, flowers, and insects.
Check Your Progress 1 ~ 1
0 Define species according to the biological species concept.
f) Describe the limitations of the biological species concept.
Categorize the different types of reproductive barriers as being either
a prezygotic or postzygotic barrier, and give an example of each.
Compare and contrast allopatric speciation with sympatric
speciation. Give an example of each.
0 Relate adaptive radiation to Darwin's finches.
Woodpecker
finch
Grasping
beaks
Warbler
finch
Probing
beaks
CHAPTER 16 Evolution on a Large Scale 2
1
Connections and Misconceptions
Are there examples of polyploid species in animal!
In general, polyploidy is rarer in
animals than in plants. However,
there are examples of polyploid
insects and fish, and polyploidy also
appears to occur frequently in the
amphibians, specifically in salaman-
ders. In 1999, scientists reported a
polyploid rat species (Typanoctomys
barrerae) in Argentina, but later genetic analysis refuted tf
claim. Most geneticists believe that polyploidy in mamm<
is unlikely due to the well-defined role of mammalian s
chromosomes and the balance between the number of autt
somes and sex chromosomes.
Connecting the Concepts
For more information on the concepts presented in this
section, refer to the following discussions.
Section 9.4 provides background information on how
nondisjunction causes changes in chromosome number.
Section 14.1 examines the role that the Galapagos finches
played in establishing natural selection as the mechanism
for evolutionary change.
Cactus
ground
finch
Figure 16.9 Darwin's finches.
Each of Darwin's finches is adapted to gathering anc
eating a different type of food. Tree finches
have beaks largely adapted to eating
insects and, at times, plants.
Sharp-beaked
ground finch
Crushing
beaks
Ground finches have beaks
adapted to eating the
flesh of the prickly-
pear cactus or
different-sizec
seeds.
272 PARTTHREE Evolution
e.
16.2 The Fossil Record
Learning Outcomes
Upon completion of this section, you should be able to
1. Describe how the geological timescale is divided into eras, periods,
and epochs.
2. Contrast the gradualistic model of evolution with the punctuated
equilibrium model of evolution.
3. Explain some of the ways in which mass extinctions of organisms
may have occurred.
The history of the origin and extinction of species on Earth is best discovered
by studying the fossil record (Fig.16.10). Fossils are the traces and remains of
past life or any other direct evidence of past life. Paleontology is the science
of discovering and studying the fossil record and, from it, making decisions
about the history of species.
The Geological Timescale
Because life-forms have evolved over time, the strata (layers of
sedimentary rock, see Fig. 14.2a) of the Earth' s crust contain
different fossils. By studying the strata and the fossils they
contain, have been able to construct a geological
timescale (Table 16.1). It divides the history of life on Earth
into eras, then periods, and then epochs and describes the
types of fossils common to each of these divisions of time.
Notice that, in the geological timescale, only the periods of
the Cenozoic era are divided into epochs, meaning that more
attention is given to the evolution of primates and flowering
plants than to the earlier evolving organisms. Despite an
epoch assigned to modern civilization, Animation
humans have only been around about r oo GeologicaiHistory
of the Earth
.04% of the history of life.
In contrast, prokaryotes existed alone for some 2 billion years
before the eukaryotic cell and multicellularity arose during Precam-
brian time. Some prokaryotes became the first photosynthesizers to
add oxygen to the atmosphere. The presence of oxygen may have
spurred the evolution of the eukaryotic cell and multicellularity
during the Precambrian. All major groups of animals evolved dur-
ing what is sometimes called the Cambrian explosion. The fossil
record for Precambrian time is meager, but the fossil record for the
Cambrian period is rich. The evolution of the invertebrate external
Figure 16.10 Fossils.
a. A fern leaf from 245 million years ago (MYA) remains because it was buried
in sediment that hardened to rock. b. This midge (40 MYA) became embedded
in amber (hardened resin from a tree). c. Most fossils, such as this early
insectivore mammal (47 MYA) are remains of hard parts because they do not
decay as the soft parts do. d. This dinosaur footprint (135 MYA) is a sign of
ancient life. e. This tree trunk (190 MYA) is petrified because minerals have
replaced the original tissues.
CHAPTER 16 Evolution on a Large Scale 2:
Table 16.1 The Geological Timescale: Major Divisions of Geological Time
and Some of the Major Evolutionary Events That Occurred
Era Period
Quaternary
Cenozoic
Tertiary
Cretaceous
Jurassic
Mesozoic
Triassic
Permian
Carboniferous
Devonian
Paleozoic
Silurian
Ordovician
Cambrian
Precambrian time
Epoch MYA Plant and Animal Life
Holocene 0-0.01 AGE OF HUMAN CIVILIZATION; Destruction of
ecosystems accelerates extinctions.
SIGNIFICANT MAMMALIAN EXTINCTION
Pleistocene 0.01 - 2
Pliocene 2-6
Miocene 6-24
Oligocene 24- 37
Eocene 37-58
Paleocene 58- 65
MASS EXTINCTION:
65- 144
144-208
MASS EXTINCTION:
208-250
Modern humans appear; modern plants spread
and diversify.
First hominids appear; modern angiosperms
flourish.
Ape-like mammals, grazing mammals, and insects
flo"'''"' 9""''"d' 'pceod; ood foce>t'
Monkey-like primates appear; modern
.,
angiosperms appear.
All modern orders of mammals are present;
subtropical forests flourish.
Primates, herbivores, carnivores, insectivores are
prese.nt;. angiosperms diversify.
50% OF ALL SPECIES, DINOSAURS, AND
Placental mammals ill"!d modern insects appear;
angiosperms spread .and conifers persist.
Dinosaurs flourish; birds and
angiosperms appear. ,
48 % OF ALL SPECIES,
First mammals and dinosaurs appear; forests
of conifers and cycads dominate land; corals
and molluscs dominate seas.
MASS EXTINCTION (THE "GREAT DYING" ) : 83% OF ALL SPECIES ON
250-286
286-360
MASS EXTINCTION:
360-408
408-438
Reptiles diversify; amphibians decline; and
gymnosperms diversify.
Amphibians diversify; reptiles appear; and
diversify. Age of great coal-forming forests.
OVER 50 % OF COASTAL MARINE SPECIES ,
Jawed fishes diversify; insects and amphibians
appear; seedless vascular plants diversify and
seed plants appear.
First jawed fishes and seedless vascular
plants appear.
MASS EXTINCTION: OVER 57 % OF MARINE SPECIES
438-510
510-543
600
1,400-700
2,000
2,500
3,500
4,500
Invertebrates spread and diversify; jawless fishes
appear; nonvascular plants appear on land.
Marine invertebrates with skeletons are dominant
and invade land, and marine algae flourish.
Oldest soft-bodied invertebrate fossils.
Protists evolve and diversify.
Oldest eukaryotic fossils.
0
2
accumulates in atmosphere.
Oldest known fossils (prokaryotes).
Earth forms.
CORALS
274 PARTTHREE Evolution
Connections and Misconceptions
What is the Burgess Shale?
The Burgess Shale is the name
for a rock formation in the Cana-
dian Rocky Mountains near the
Burgess Pass. Around 525 MYA,
this region was located along
the coast. It is believed that an
earthquake caused a landslide that almost instantly buried
much of the marine life living in the shallow coastal waters.
Unlike many fossil beds, the Burgess Shale contains the
remains of soft-shelled organisms, such as worms and sea
cucumbers, as well as other organisms from the Cambrian
explosion-a period of rapid diversification in marine life
around 545 MYA. Over 60,000 unique types of fossils have
been found in the Burgess Shale (including the trilobite fossil
shown here), making this fossil bed one of our most valuable
assets for studying the early evolution of life in the oceans.
Figure 16.11 Paceofevolution.
a. According to the gradualistic model, new
species evolve slowly from an ancestral species.
b. According to the punctuated equilibrium
model, new species appear suddenly and then
remain largely unchanged until they become
extinct.
a. Gradualistic model
skeleton accounts for this increase in the number of fossils. Perhaps this skeleton.
which impedes the uptake of oxygen, couldn't evolve until oxygen was plentiful.
Or perhaps the external skeleton was merely a defense against predation.
The origin of life on land is another interesting topic. During the Paleozoic
era, plants were present on land before animals. Nonvascular plants preceded
vascular plants, and among these, cone-bearing plants (gymnosperms) preceded
flowering plants (angiosperms). Among vertebrates, the fishes were aquatic, and
the amphibians invaded land. The reptiles, including dinosaurs and birds, shared
an amniote ancestor with the mammals. The number of species in the world has
continued to increase until the pre ent time, despite the occurrence of five mas1
extinctions, including one significant mammalian extinction during the histof)
of life on Earth.
The Pace of Speciation
Darwin theorized that evolutionary changes occur gradually. In other words, he sup-
ported a gradualistic model to explain the pace of evolution. Speciation probably
occurs after populations become isolated, with each group continuing slowly on
its own evolutionary pathway. The gradualism model often shows the evolutionaf)
history of groups of organisms by drawing the type of evolutionary tree shown in
Figure 16.11a. In this diagram, note that an ancestral species has given rise to two
separate species, represented by a slow change in plumage color. The gradualistic
model suggests that it is difficult to indicate when speciation has occurred because
there would be so many transitional links. In some cases, it has been possible to
trace the evolution of a group of organisms by finding transitional links.
More often, however, species ap(.>ear quite suddenly in the fossil record, and
then they remain essentially unchan'ged phenotypically until they undergo extinc
tion. Some paleontologists have therefore developed a punctuated equilibrium
model to explain the pace of evolution. The model says that a period of equilib-
Time
Change
Time
Change
b. Punctuated equilibrium model
I,
I.
c
d
j
j
j
s
rium (no change) is punctuated (interrupted) by speciation. Figure l6.llb shows
the type of diagram paleontologists prefer to use when representing the history of
evolution over time. This model suggests that transitional links are less likely to
become fossils and less likely to be found. Speciation most likely involves only an
isolated subpopulation at one locale. Only when this new subpopulation expands
and replaces existing species is it apt to show up in the fossil record.
The differences between these two models are subtle, especially when
we consider that the "sudden" appearance of a new specie in the fossil record
could represent many thousands of years because geological time is measured
in millions of years.
Mass Extinctions of Species
As researchers have noted, most species exist for only a limited amount of time
(measured in millions of years), and then they die out (become extinct). Mass
extinctions are the disappearance of a large number of species or a higher taxo-
nomic group within a relatively short period of time. The geological timescale
in Table 16.1 shows the occurrence of five mass extinctions: at the ends of the
Ordovician, Devonian, Permian, Triassic, and Cretaceous periods. Also, there
was a significant mammalian extinction at the end of the Pleistocene epoch.
While many factors contribute to mass extinctions, two possible types of events
(continental drift and meteorite impacts) have particular significance.
Continental drift-the movement of continents-has contributed to sev-
eral extinctions. You may have noticed that the coastlines of several continents
are mirror images of each other. For example, the outline of the west coast of
Africa matches that of the east coast of South America. Also, the same geological
structures are found in many of the areas where the continents touched at one
time. A single mountain range runs through South America, Antarctica, and
Australia, for example. But the mountain range is no longer continuous because
the continents have drifted apart. The reason the continents drift is explained by
a branch of geology known as plate tectonics, which is based on the fact that the
Earth's crust is fragmented into slab-like plates that float on a lower, hot mantle
layer (Fig. 16.12). The continents and the ocean basins are a part of these rigid
plates, which move like conveyor belts.
Continental plates meet
along a fault line, shift,
and cause earthquakes.
Continental plate
is folded into
Earth's
CHAPTER 16 Evolution on a Large Scale
Figure 16.12 Plate tectonics.
The Earth's surface is divided into several solid tectonic plates flo;
on the fluid magma beneath them. At rifts in the ocean floor, two
gradually separate as fresh magma wells up and cools, enlarging j
plates. Mountains, including volcanoes, are raised where one plat
pushes beneath another at subduction zones. Where two plates s
grind past each other at a fault line, tension builds up, which is rel1
occasionally in the form of eathquakes.
Oceanic plate sinks beneath
continental plate and melts
into magma again.

_ Hot magma rises

to the surface
and cools.
276 PART THREE Evolution
a.
Pangaea:
North
America
-o
"'
Eurasia
South 1- Africa
America Q -1
~ {
India
Antarctica
Late Paleozoic, 250 MYA
...
'\
Equator
Australia
North
America
...
Eurasia
b.
'
South
America
'
Most modern continents
had formed by the end of
the Mesozoic, 65 MYA
I
'
Africa
I
.if"
Jl' Antarctica
Figure 16.13 Continental drift.
t
f
Equator
India
t
Australia
a. About 250 MYA, all the continents were joined into a supercontinent
called Pangaea. b. By 65 MYA, all the continents had begun to separate.
This process is continuing today. North America and Europe are
separating at a rate of about 2 centimeters per year.
Connecting the Concepts
For more information on the material in this section, refer to
the following discussions.
Section 14.2 describes how the fossil record is used as
evidence of evolutionary change.
Sections 18.1 and 19.1 provide an overview of the
evolution ofthe plants and animals, respectively.
Section 32.1 examines some of the influences that humans
are having on the diversity of life.
The loss of habitat is a significant cause of extinctions, and continental
drift can lead to massive habitat changes. We know that 250 million years ago
(MYA), at the time of the Permian mass extinction, all the Earth's continents
came together to form one supercontinent called Pangaea (Fig. 16.13a). The
result was dramatic environmental change through the shifting of wind patterns,
ocean currents, and most importantly the amount of available shallow marine
habitat. Marine organisms suffered as the oceans merged, and the amount of
shoreline, where many marine organisms lived, was drastically reduced. Spe-
cies diversity did not recover until some continents drifted away from the poles,
shorelines increased, and warmth returned (Fig. 16.13b ). Terrestrial organisms
were affected as well because the amount of interior land, which tends to have
a drier and more erratic climate, increased. Immense glaciers developing at
the poles withdrew water from the oceans and chilled even once tropical land.
The other event that is known to have contributed to mass extinctions
is the impact of a meteorite as it crashed into the Earth. The result of a large
meteorite striking Earth could have been similar to that of a worldwide atomic
bomb explosion: A cloud of dust would have mushroomed into the atmosphere,
blocking out the sun and causing plants to freeze and die. This type of event
has been proposed as a primary cause of the Cretaceous extinction that saw the
demise of the dinosaurs. Cretaceous clay contains an abnormally high level of
iridium, an element that is rare in the Earth's crust but more common in mete-
orites. A layer of soot has been identified iri the strata alongside the iridium,
and a huge crater that could have been caused by a meteorite was found in the
Caribbean-Gulf of Mexico region on the Yucatan peninsula .
Connections and Misconceptions
What is the "Sixth Mass Extinction Event"?
Many ecologists now support the concept that
we are currently involved in the Earth's sixth mass
extinction event. However, unlike the first five
major events, this one is caused not by geologi-
cal or astronomical events but by human actions.
Pollution, land use, invasive species, and global
climate change associated with the burning of fossil fuels are all recognized
as contributing factors. The exact rate of species loss can be difficult to
determine, but international agencies report that the current loss of species
is between 100 and 1,000 times faster than the pre-human rates recorded by
the fossil record.
Check Your Progress-16.2
0 Summarize how the fossil record is the best evidence for
macroevolution.
Explain the phrase "punctuated equilibrium model."
Relate the mass extinctions of species with the types of events that
can cause them.
Discuss some of the major evolutionary events that occurred during
different periods of geological time to form the geological timescale
(Table 16.1).
16.3 Systematics
Learning Outcomes
Upon completion of this section, you should be able to
1. List the hierarchical levels of Linnaean classification from the
most inclusive to the least inclusive and explain how this type of
classification is useful to biologists.
2. Explain what information can be learned from a phylogenetic tree and
list some of the types of information that is used in constructing them.
3. Contrast a homologous structure with an analogous structure.
4. Define cladistics and explain how this method may be used to study
the evolutionary relationships between groups of organisms.
5. List the three domains of living organisms, and describe the general
characteristics of organisms contained within each domain.
I field of biology, but especially systematics, are dedicated to
mlanding the evolutionary history of life on Earth. Systemat-
1\ very analytical and relies on a combination of data from the
sil record and comparative anatomy and development, with
emphasis today on molecular data, to determine phylogeny,
Domain
Bacteria
evolutionary history of a group of organisms. Classification is a part of
tematics because ideally organisms are classified according to our present
mtanding of evolutionary relationships.
nnaean Classification
Taxonomy is the branch of biology concerned with identifying, nam-
. and classifying organisms. A taxon (pl., taxa) is an organism or a
poforganisms at a particular level in a classification system. The
mial system of nomenclature assigns a two-part name to each
of organism. For example, the plant in Figure 16.14 has been
ed Cypripedium acaule. This name means that the plant is
the genus Cypripedium and that the specific epithet is acaule.
,e that the scientific name is in italics and only the genus is
nalized. The genus can be abbreviated to a single letter if the
name has been given previously and if it is used with a specific
. For example, C. acaule is an acceptable way to designate
plant. The name of an organism usually tells you something about
. In this instance, the genus name, Cypripedium, refers to the
of the flower, and the specific epithet, acaule, says that the flower
independent stem.
Why do organisms need scientific names? And why do scientists use Latin,
common names, to describe organisms? There are several reasons. First,
name will vary from country to country because clifferent countries use
languages. Second, even people who speak the same language sometimes
ilifferent common names to describe the same organism-for example, bowfin,
16.14 Taxonomy hierarchy.
CHAPTER 16 Evolution on a Large Scale 277
Kingdom
Plantae
Monocotyledones
Family
Asparagales
Orchidaceae
Species
Cypripedium
Cypripedium
acaule
is the most inclusive of the classification categories. The plant kingdom is in the domain Eukarya.ln the plant kingdom are several phyla, each represented here by
I The phylum Anthophyta has only two classes (the monocots and eudicots). The class Monocotyledones encompasses many orders. In the order Orchid ales
lllln,,f,miilio< in the family Orchidaceae are many genera; and in the genus Cypripedium are many species-for example, Cypripedium acaule. (This illustration is
i and doesn't necessarily show the correct number of subcategories.)
278 PART THREE Evolution
Ovis aries
(sheep)
Ovis
Bos taurus
(cattle)
Cervus elaphus
(red deer)
Figure 16.15 Classification and phylogeny.
The classification and phylogenetic tree for a group of organisms is
ideally constructed to reflect their phylogenetic history. A species is most
closely related to other species in the same genus, more distantly related
to species in other genera of the same family, and so forth on through
order, class, phylum, and kingdom.
Rangifer tarandus
(reindeer)
Species
Genus
Family
Order
grindle, choupique, and cypress trout are all the same
common fish, Amia calva. Furthermore, between
countries the same common name is sometimes given
to different organisms. A robin in England is very dif
ferent from a robin in the United States, for example.
Latin, on the other hand, is a universal language that not
too long ago was well known by most scholars, many
of whom were physicians or clerics. When scientists
throughout the world use a scientific binomial name.
they know they are speaking of the same organism.
Today, taxonomists use several categories of
clas ification created by Swedish biologist Carl Lin-
naeus in the 1700s to show varying levels of similar-
ity: species, genus, family, order, class, phylum,
and kingdom. More recently, a higher taxonomic cat-
egory, the domain, has been added to this list. There
can be several species within a genus, several genera
within a family, and so forth. In this hierarchy, the
higher the category, the more inclusive it is (Fig. 16.14).
Therefore species in the same domain have general traits
in common, while those in the same genus have quite specific traits in common.
In most cases, each category of classification can be subdivided into
three additional categories, as in superorder, order, suborder, a.nd infraorder.
Considering these, there are more than 30 categories of classification.
Phylogenetic Trees
Figure 16.15 shows how the classification of groups of organisms allows us
to construct a phylogenetic tree, a diagram that indicates common ancestors
and lines of descent (lineages). The common ancestor at the base of the tree
has traits that are shared by all the other groups in the tree. For example, the
Artiodactyla are characterized by having hoofs with an even number of toes. On
the other hand, notice that the Cervidae have antlers but the Bovidae have no
antlers. Finally, among the Cervidae, the antlers are highly branched in the red
deer but palmate (having the shape of a hand) in reindeer. As the lineage moves
from common ancestor to common ancestor, the traits become more pecific to
just particular groups of animals. It is this progression in
specificity that allows classification categories to serve as a
basis for depicting a phylogenetic tree.
Tracing Phylogeny
Animation
Phylogenetic
Trees
While Figure 16.15 makes use of morphological data, systematists today use
a multitude of data in order to discover the evolutionary relationships between
species. They rely heavily on a combination of fossil record data, morpho-
logical data, and molecular data to determine the correct sequence of common
ancestors in any group of organisms.
Morphological data include homologies, which are similarities among
organisms that stem from having a common ancestor. Comparative anatomy,
including embryological evidence and fossil data, provides information regard-
ing homology. Homologous structures are related to each other through com-
mon descent. The forelimbs of vertebrates are homologous because they contain
the same bones organized in the same general way as in a common ancestor (see
Fig. 14.15). This is the case even though a horse has but a single digit and toe (the
hoof), while a bat has four lengthened digits that support its membranous wings.
Deciphering homology is sometimes difficult because of convergent
evolution. Convergent evolution is the acquisition of the same or similar traits
in distantly related lines of descent. Similarity due to convergence is termed
analogy. The wings of an insect and the wings of a bat are analogous. You may
recall from Chapter 14 that analogous structures have the same function in
different groups but organisms with these structures do not have a recent com-
mon ancestor. Analogous structures arise because of adaptations to the same
~ p of environment. Both cactuses and spurge are adapted similarly to a hot,
dry environment, and both are succulent, spiny, flowering plants. However, the
details of their flower structure indicate that these plants are not closely related.
Speciation occurs when mutations bring about changes in the base-pair
sequences of DNA. Systematists, therefore, assume that, the more closely spe-
cies are related, the fewer changes will be found in DNA base-pair sequences.
Because molecular data are straightforward and numerical, they can sometimes
sort out relationships obscured by inconsequential anatomical variations or
convergence. Computer software breakthroughs have made it possible to
analyze nucleotide sequences quickly and accurately. Also, these analyses are
available to anyone doing comparative studies through the Internet, so each
mvestigator doesn't have to start from scratch. The combination of accuracy
and availability of vast amounts of data, even entire genomes, has made
molecular systematics a standard way to study the relatedness of organisms.
All cells have ribosomes essential for protein synthesis, and the gene
!hat code for ribosomal RNA (rRNA) have changed very slowly during evolu-
tion because drastic changes lead to malfunctioning cells. Therefore, compara-
uve rR A sequencing provides a reliable indicator of the similarity between
organisms. Ribosomal RNA sequencing helped investigators conclude that all
tving things can be divided into the three domains.
One study involving DNA differences produced the data shown in
Figure 16.16. Notice the close relationship between chimpanzees and humans.
0
Galago Capuchin Green monkey Rhesus monkey
~
10

0
Cl
<t
VI
20
:v
~
.....
0
VI 30
c:
~
:ll
40
I
I
CHAPTER 16 Evolution on a Large Scale 279
Gibbon Chimpanzee Human
I I
I
I
hips of certain primate species are based on a study of their genomes. The length of the branches indicates the relative number of DNA base-pair differences
the groups. These data, along with knowledge of the fossil record for one divergence, make it possible to suggest a date for the other divergences in the tree.
280 PARTTHREE Evolution
03
Qi
Q)
"E ()


c
Qi
"' "' .!!!
Q) c
Q) c
"'
Notochord in embryo
Vertebrae
Lungs
Three-chambered heart
Internal fertilization
Amniotic membrane in egg
Four bony limbs
Long cylindrical body
a.
b.
Figure 16.17 Constructing a cladogram.
a. First, a table is drawn up, listing characters for all the taxa. An
examination of the table shows which characters are ancestral
(notochord, aqua) and which are derived (lavender, orange, and yellow).
The shared derived characters distinguish the taxa. b. In a cladogram,
the shared derived characters are sequenced in the order they evolved
and are used to define clades. A clade contains a common ancestor and
all the species that share the same derived characters (homologies). Four
bony limbs and a long, cylindrical body were not used in constructing
the cladogram because they are in scattered taxa.
This relationship has recently been recognized by the designation of a new
subfamily, Homininae, that includes not only chimpanzees and humans but
also gorilla . Molecular data indicate that gorillas and chimpanzees are more
closely related to humans than they are to orangutans. Below the taxon subfam-
ily, humans and chimpanzees are placed together in their own tribe, a rarely
used classification category.
Cladistics and Cladograms
Cladistics is a way to trace the evolutionary history of a group by using
shared traits derived from a common ancestor to determine relationships.
These traits are then used to construct phylogenetic trees called cladograms.
A cladogram depicts the evolutionary hi tory of a group based on the avail-
able data.
The first step in constructing a cladogram is to draw up a table that
summarizes the traits of the species being compared. At least one but prefer-
ably several species are considered an outgroup. The outgroup is oot part of
the study group, also called the ingroup. In Figure 16.17a, lancelets are the
outgroup because, unlike the pecies in the ingroup, they are not vertebrate .
Any trait, such as a notochord, found in both the outgroup and the in group is a
shared ancestral trait, presumed to have been present in a common ancestor to
both the outgroup and ingroup. Ancestral traits are not shared derived traits and
therefore are not used to construct a cladogram. They merely help us determine
which traits will be used to construct the cladogram.
A rule that many cladists follow is the principle of parsimony, which
states that the least number of assumptions is the most probable. Thus, they
construct a cladogram that minimizes the number of assumed evolutionary
changes or that leaves the fewest number of derived traits unexplained.
Therefore, any trait in the table found in scattered species (in this case.
four bony limbs and a long, cylindrical body) is not used to construct the
cladogram because we would have to assume that these traits evolved more
than once among the species of the study group. The other difference are
designated as shared derived traits-that is, they are homologies shared
by only certain species of the study group. Combining the data regarding
shared derived traits will tell us how the members of the ingroup are related
to one another.
The Cladogram
A cladogram contains everal clades; each clade includes a common ancestor
(at the circles) and all of its descendant species. The cladogram in Figure
16. l7b has three clades (l-3), which differ in size because the first includes
the other two and so forth. All the species in the study group belong to a clade
that has vertebrae; only newts, snakes, and lizards are in a clade that has lungs
and a three-chambered heart; and only snakes and lizards are in a clade that
has an amniotic egg and internal fertilization. (An amniotic egg has a sac thai
surrounds and protects the embryo-fish and amphibian eggs do not have this
sac.) Following the principle of parsimony, this is the sequence in which these
traits must have evolved during the evolutionary history of vertebrates. Any
other arrangement of species would produce a less parsimonious evolutionary
sequence-that is, a tree that would be more complicated.
A cladogram is objective-it lists the traits used to construct the
cladogram. Cladists typically use much morphological, fossil, and molecu-
lar data to construct a cladogram. Still, cladists regard a cladogram as a
hypothesis. Whether our tree is consistent with the one, true evolutionary
history of life can be tested, and modifications can be made on the basis of
additional data.
Linnaean Classification Versus Cladistics
Figure 16.18 illustrates the types of problems that arise when trying to
reconcile Linnaean classification with the principles of cladistics. Figure
16.18, which is based on cladistics, shows that birds are in a clade with
crocodiles, with which they share a recent common ancestor. This ancestor
had a gizzard. An examination of the skulls of crocodiles and birds would
show other derived traits that they share. Birds have scaly skin and share
this ancestor with other reptiles as well. However, Linnaean classification
places birds in their own group, separate from crocodiles and from reptiles
in general. In many other instances, Linnaean classification is not consis-
tent with new understandings about phylogenetic relationships. Therefore,
some cladists have proposed a different system of classification, called the
International Code of Phylogenetic Nomenclature, or PhyloCode, which
sets forth rules for the naming of clades. Other biologists are hoping to
modify Linnaean classification to be consistent with the principles of
cladistics.
Two major problems may be unsolvable: ( I) Clades are hierarchical, as
are Linnaean categories. However, there may be more clades than Linnaean
laxonomic categories, and it is therefore difficult to equate clades with taxons.
(2) The taxons are not necessarily equivalent in the Linnaean system. For
example, the family taxon within Kingdom Plantae may not be equivalent
lo the family taxon in Kingdom Animalia. Because of such problems, some
cladists recommend abandoning Linnaeus altogether .

mammals turtles
Figure 16.18 Cladistic classification.
CHAPTER 16 Evolution on a Large Scale 21
egg,
internal fertilization
Taxonomic designations are based on evolutionary history. Each taxon includes a common ancestor
and all of its descendants.
282 PARTTHREE Evolution
Figure 16.19 Three-do main system.
In this system, the prokaryotes are in the domains Bacteria and
Archaea. The eukaryotes are in the domain Eukarya, which contains
four kingdoms for the protists, animals, fungi, and plants.
cyanobacteria
Bacteria
Prokaryotic, unicellular
organisms
Lack a membrane-bounded
nucleus
Reproduce asexually
Heterotrophic by absorption
Autotrophic by
chemosynthesis or by
photosynthesis
Move by flagell a
common ancestor
The Three-Domain System
Classification systems change over time. Historically, most biologi sts utilized
the five-kingdom system of classification, which contains kingdoms for the
plants, animals, fungi, protists, and monerans. Organisms were placed into these
kingdoms based on type of cell (prokaryotic or eukaryotic), level of organization
(unicellular or multicellular), and type of nutrition. In the five-kingdom system,
the monerans were distinguished by their structure-they were prokaryotic (lack
a membrane-bounded nucleus)-whereas the organisms in the other kingdoms
were eukaryotic (have a membrane-bounded nucleu ). The kingdom Monera
contained all prokaryotes, which evolved first, according to the fossil record.
eukaryotes
plants
animals
Eukarya
Eukaryotic, unicellular to
multicellular organisms
Membrane-bounded
nucleus
Sexual reproduction
Phenotypes and nutrition
are diverse
Each kingdom has
specializations
Flagella, if present, have a
9 + 2 organizati on
prokaryotes
proti sts
Archaea
Prokaryotic, unicell ular
organisms
Lack a membrane-bounded
nucleus
Reproduce asexuall y
Many are autotrophic by
chemosynthesis; some are
heterotrophic by absorption
Unique rRNA base
sequence
Distinctive plasma
membrane and cell wall
chemistry
Sequencing the genes for rRNA has provided new information that calls
into question the five-kingdom system of classification. Aside from molecular
data, cellular data also suggest that there are two groups of prokaryotes, named
the Bacteria and the Archaea. These two groups are so fundamentally different
from each other they have been assigned to separate domains, a category of
classification that is higher than the kingdom category. The Bacteria arose first,
followed by the Archaea and then the Eukarya (Fig. 16.19). The Archaea and
Eukarya are more closely related to each other than either is to the Bacteria.
Systematists, using the three-domain system of classification, are in the process
of sorting out what kingdoms belong within domain Bacteria and domain
Archaea. Domain Eukarya contains kingdoms for protists, animals, fungi,
and plants. Later in this text, we will study the individual kingdoms that occur
v;ithin the domain Eukarya. The protists do not share one common ancestor, and
some suggest that the kingdom should be divided into many different kingdoms.
The number of kingdoms is still being determined among
systematists, illustrating that classification can be changed
as new data become available.
Connecting the Concepts
Animation
For more information on the topics presented in this section, refer to the
following discussions.
Section 14.2 provides an overview of how comparative anatomy is used to
study evolution.
Section 17.3 examines the evolution of the prokaryotic domains of life.
Figure 19.4 illustrates the phylogenetic tree of animal evolution.
Check Your Progress 16.3
0 List the categories of classification in order from smallest to largest.
G Name the types of data used to determine evolutionary
relationships.
Contrast homologous structure with analogous structure.
Explain why the sequencing of ribosomal RNA (rRNA) is done for
evolutionary studies.
0 Explain why the Linnaean classification is difficult to reconcile with
the principles of cladistics.
0 Explain the following: If Band Dare fishes, what other animal
(designated by a letter) in this cladogram is also a fish? Explain.
A B c 0
CHAPTER 16 Evolution on a Large Scale
- -------------
284 PART THREE Evolution
Media Study Tools
www.mhhe.com/maderessentials3
~ ~ c o n n e c t ~ e
I BIOLOGY
Enhance your study of this chapter with study tools and practice tests. Also ask your instructor
about the resources available through ConnectPius, including the media-rich eBook, interactive
learning tools, and animations.
The Chapter in Review
Summary
16.1 Speciation and Macroevolution
Macroevolution is evolution on a large scale because it considers the
history of life on Earth. Macroevolution begins with speciation, the origin
of new species. Without speciation and the extinction of species, life on
Earth would not have a history.
Defining Species
The biological concept of species
recognizes a species by its inability to produce viable fertile offspring
with members of another group.
is useful because species can look similar, and members of the same
species can have different appearances.
has its limitations. Hybridization does occur between some
species, and it is only relevant to extant (living or not extinct) sexually
reproducing organisms.
Reproductive Barriers
Prezygotic and postzygotic barriers keep species from reproducing with
one another.
Prezygotic isolating mechanisms involve habitat isolation, temporal
isolation, and behavioral isolation.
Postzygotic isolating mechanisms prevent hybrid offspring from
developing or breeding if reproduction has been successful.
Models of Speciation
Allopatric speciation and sympatric speciation are two models of speciation.
The allopatric speciation model proposes that a geographic barrier
keeps groups of populations apart. Meanwhile, prezygotic
and postzygotic isolating mechanisms develop, and these
prevent successful reproduction if these two groups come in
future contact.
The sympatric speciation model proposes that a geographic barrier is
not required for speciation to occur.
16.2 The Fossil Record
The fossil record, as outlined by the geological timescale, traces the
history of life in broad terms. It has been possible to absolutely date fossils
by using radioactive dating techniques.
The fossil record
can be used to support a gradualistic model : Two groups of
organisms arise from an ancestral species and gradually become two
different species.
also supports the punctuated equilibrium model : A period of
equilibrium (no change) is interrupted by speciation within a
relatively short period of time.
shows that at least five mass extinctions, including one significant
mammalian extinction, have occurred during the history of life on
Earth. Two major contributors to mass extinctions are the loss of
habitat due to continental drift and the disastrous results from a
meteorite impacting Earth.
16.3 Systematics
Systematics is the study of the evolutionary relationships among
all organisms, past and present. Systematics includes
classification. In the Linnaean system of classification, every
organism is assigned a scientific name, which indicates its genus
and specific epithet. Species are also assigned to a family, order,
class, phylum, kingdom, and domain according to their molecular
and structural similarities as well as evolutionary relationships to
other species.
Phylogeny depicts the evolutionary history of a group of organisms.
Systematics relies on the fossil record, homology, and molecular data
to determine relationships among organisms.
Cladists use shared derived characters to construct cladograms. In a
cladogram, a clade consisting of a common ancestor and all the
species derived from that ancestor, the species have shared derived
characteristics.
Linnaean classification has come under severe criticism because it
does not always follow the principles of cladistics in the grouping of
organisms.
Classification Systems
The three-domain system uses molecular data to designate three
evolutionary domains: Bacteria, Archaea, and Eukarya:
~
Bactena
Domains Bacteria and Archaea contain prokaryotes.
Domain Eukarya contains kingdoms for the protists, animals, fungi,
and plants.
Key Terms
adaptive radiation 271
allopatric speciation 270
analogous structure 279
analogy 279
clade 280
cladistics 280
cladogram 280
class 278
common ancestor 268
convergent evolution 279
domain 278
domain Archaea 283
domain Bacteria 283
domain Eukarya 283
evolutionary tree 274
family 278
five-kingdom system 282
fossil 272
genus 278
homologous structure 278
kingdom 278
macroevolution 266
mass extinction 275
order 278
paleontology 272
phylogenetic tree 278
phylogeny 277
phylum 278
postzygotic isolating
mechanism 269
prezygotic isolating
mechanism 268
speciation 266
species 278
sympatric speciation 270
systematics 277
taxon (pl., taxa) 277
taxonomy 277
three-domain system 283
Testing Yourself
Choose the best answer for each question.
1. A biological species
a. always looks different from other species.
b. always has a different chromosome number from that of
other species.
c. is reproductively isolated from other species.
d. never occupies the same niche as other species.
For questions 2-9, indicate the type of isolating mechanism described in
each scenario.
Key:
a. habitat isolation e. gamete isolation
b. temporal isolation f. zygote mortality
c. behavioral isolation g. hybrid sterility
d. mechanical isolation h. low F
2
fitness
2. Females of one species do not recognize the courtship behaviors of
males of another species.
3. One species reproduces at a different time of year than another
species.
4. A cross between two species produces a zygote that always dies.
5. Two species do not interbreed because they occupy different
areas.
6. A hybrid between two species produces gametes that are not
viable.
7. Two species of plants do not hybridize because they are visited by
different pollinators.
B. The sperm of one species cannot survive in the reproductive tract of
another species.
9. The offspring of two hybrid individuals exhibit poor vigor.
CHAPTER 16 Evolution on a Large Scale 21
10. Complete the following diagram illustrating allopatric speciation
by using these phrases: genetic changes (used twice), geographic
barrier, species 1, species 2, species 3.
a.
11. Transitional links are least likely to be found if evolution proceeds
according to the
a. gradualistic model.
b. punctuated equilibrium model.
12. Which of the following is the scientific name of an organism?
a. Rosa rugosa d. rugosa rugosa
b. Rosa e. Both a and d are correct.
c. rugosa
13. Which of these statements best pertains to taxonomy?
a. Species always have three-part names, such as Homo
sapiens sapiens.
b. Species are always reproductively isolated from other species.
c. Species share ancestral traits but may have their own unique
derived traits.
d. Species always look exactly alike.
e. Both c and dare correct.
14. Which of the following groups are domains? Choose more than one
answer if correct.
a. bacteria d. animals
b. archaea e. plants
c. eukarya
15. Which pair is mismatched?
a. homology-character similarity due to a common ancestor
b. molecular data-DNA strands match
c. fossil record- bones and teeth
d. homology-functions always differ
e. molecular data- DNA and RNA data
16. One benefit of the fossil record is
a. that hard parts are more likely to fossilize.
b. that fossils can be dated.
c. its completeness.
d. that fossils congregate in one place.
e. All of these are correct.
17. The discovery of common ancestors in the fossil record, the
presence of homologies, and molecular data similarities help
scientists determine
a. how to classify organisms.
b. the proper cladogram.
c. how to construct phylogenetic trees.
d. how evolution occurred.
e. All of these are correct.
286 PARTTHREE Evolution
18. In cladistics,
a. a clade must contain the common ancestor plus all its
descendants.
b. shared derived characters help construct cladograms.
c. the principle of parsimony states that the simpler hypothesis is
preferred.
d. the species in a clade share homologous structures.
e. All of these are correct.
19. Answer these questions about the following cladogram.
amniotic egg,
internal fertilization
lungs,
three-chambered heart
a. This cladogram contains how many clades? How are they
designated in the diagram?
b. What character is shared by all animals in the study group? What
characters are shared by only snakes and lizards?
c. Which animals share the most recent common ancestor? How
do you know?
20. Allopatric, but not sympatric, speciation requires
a. reproductive isolation.
b. geographic isolation.
c. spontaneous differences in males and females.
d. prior hybridization.
e. rapid rate of mutation.
21. Which kingdom is mismatched?
a. Protista- domain Bacteria
b. Protista- single-celled algae
c. Plantae-flowers and mosses
d. Animalia- arthropods and humans
e. Fungi- molds and mushrooms
22. Many new species evolving in various environments from a
common ancestor is called
a. cladistics.
b. the gradualistic model of evolution.
c. adaptive radiation.
d. convergent evolution.
e. the PhyloCode.
Thinking Scientifically
1. Using as many terms as necessary (from both X andY axes), fill in
the proposed phylogenetic tree for vascular plants.
Ferns Conifers Ginkgos Monocots Eud1cots
vascular tissue X X X X X
produce seeds X X X X
naked seeds X X
needle-like leaves X
fan-shaped leaves X
enclosed seeds X X
one embryonic leaf X
two embryonic leaves X
a. d. e. g. h.
vascular tissue
2. The Hawaiian Islands are located thousands of kilometers from any
mainland. Each island arose from the sea bottom and was colonized
by plants and animals that drifted in on ocean currents or winds.
Each island has a unique environment in which its inhabitants have
evolved. Consequently, most of the plant and animal species on the
islands do not exist anywhere else in the world.
In contrast, on the islands of the Florida Keys, there are no unique
or indigenous species. All of the species on those islands also exist
on the mainland. Suggest an explanation for these two different
patterns of speciation.
Bioethicallssue
Funding for Phylogenies
Reconstructing evolutionary relationships can have its benefits. For
example, an emerging virus is apt to jump to a related species rather than
to an unrelated species. We now know, for example, that the HIV virus
jumped from the chimpanzee-with which we share 95% of our DNA
sequence-to humans. Chimpanzees don't become ill from the virus;
therefore, studying their immune system might help us develop strategies
to combat AIDS in humans.
Cladists want to acquire more information about how genetic and
environmental factors influence bone development. Cladograms based
on bone shape and arrangement do not match up well with those based
on DNA base-pair sequencing. If we knew more about bone development
under different environmental conditions, we might be able to discover
the reason they don't match and develop evolutionary (phylogenetic)
trees in which all have confidence. To acquire the necessary data could
cost millions of dollars in research funding.
Should the public be willing to fund all types of research or
only research that has immediate medical benefits, as was described for
HIV research? Would you be willing to fund research that helps us
understand our evolutionary past, even if the medical benefit is not
immediately known? Do you think public education in cladistic research
and its benefits would be beneficial?