CAMPBELL BIOLOGY IN FOCUS

Urry • Cain • Wasserman • Minorsky • Jackson • Reece

23
Broad Patterns
of Evolution

Lecture Presentations by
Kathleen Fitzpatrick and Nicole Tunbridge

© 2014 Pearson Education, Inc.

Overview: Lost Worlds

 Past organisms were very different from those now

alive
 The fossil record shows evidence of macroevolution,
broad changes above the species level; for example
 The emergence of terrestrial vertebrates
 The impact of mass extinctions
 The origin of flight in birds

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 Figure 23.1

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 Figure 23.UN01

Cryolophosaurus skull

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Concept 23.1: The fossil record documents life’s
history
 The fossil record reveals changes in the history of
life on Earth

Video: Grand Canyon

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 Figure 23.2
1m

100
mya Rhomaleosaurus
victor
175
0.5 m 200
Dimetrodon

270
300 Tiktaalik

4.5 cm
Coccosteus cuspidatus 375
400 1 cm
Hallucigenia

500
510 2.5 cm Dickinsonia
costata
560
Stromatolites 600
1,500
3,500 Tappania
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 Figure 23.2a

Stromatolite cross
section

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 Figure 23.2b

Stromatolites

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 Figure 23.2c

Tappania

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 Figure 23.2d

2.5 cm

Dickinsonia costata

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 Figure 23.2e

1 cm
Hallucigenia

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 Figure 23.2f

4.5 cm
Coccosteus cuspidatus

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 Figure 23.2g

Tiktaalik

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 Figure 23.2h

0.5 m
Dimetrodon

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 Figure 23.2i

1m
Rhomaleosaurus victor

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The Fossil Record

 Sedimentary rocks are deposited into layers called

strata and are the richest source of fossils
 The fossil record indicates that there have been
great changes in the kinds of organisms on Earth at
different points in time

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 Few individuals have fossilized, and even fewer
have been discovered
 The fossil record is biased in favor of species that
 Existed for a long time
 Were abundant and widespread
 Had hard parts

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How Rocks and Fossils Are Dated

 Sedimentary strata reveal the relative ages of fossils

 The absolute ages of fossils can be determined by
radiometric dating
 A “parent” isotope decays to a “daughter” isotope at
a constant rate
 Each isotope has a known half-life, the time
required for half the parent isotope to decay

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 Figure 23.3

isotope remaining
Fraction of parent

Accumulating
“daughter”
isotope
½

Remaining ¼
“parent”
isotope ⅛ 1 16

1 2 3 4
Time (half-lives)
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 Radiocarbon dating can be used to date fossils up
to 75,000 years old
 For older fossils, some isotopes can be used to date
volcanic rock layers above and below the fossil

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The Geologic Record

 The geologic record is a standard time scale

dividing Earth’s history into the Hadean, Archaean,
Proterozoic, and Phanerozoic eons
 The Phanerozoic encompasses most of the time
that animals have existed on Earth
 The Phanerozoic is divided into three eras: the
Paleozoic, Mesozoic, and Cenozoic
 Major boundaries between geological divisions
correspond to extinction events in the fossil record

Animation: The Geologic Record

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 Table 23.1

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 Table 23.1a

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 Table 23.1b

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 The oldest known fossils are stromatolites, rocks
formed by the accumulation of sedimentary layers
on bacterial mats
 Stromatolites date back 3.5 billion years ago
 Prokaryotes were Earth’s sole inhabitants for more
than 1.5 billion years

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 Early prokaryotes released oxygen into the
atmosphere through the process of photosynthesis
 The increase in atmospheric oxygen that began 2.4
billion years ago led to the extinction of many
organisms
 The eukaryotes flourished in the oxygen-rich
atmosphere and gave rise to multicellular organisms

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The Origin of New Groups of Organisms

 Mammals belong to the group of animals called

tetrapods
 The evolution of unique mammalian features can be
traced through gradual changes over time

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 Figure 23.4
Reptiles Key to skull bones
(including Articular Dentary
OTHER dinosaurs and birds)
Quadrate Squamosal
TETRA-
PODS †Dimetrodon

Synapsids Early cynodont (260 mya)

Therapsids
Cynodonts
†Verylate (non-
Temporal
mammalian)
fenestra
cynodonts
(partial view)
Mammals
Hinge
Synapsid (300 mya)

Later cynodont (220 mya)

Temporal
fenestra

Hinge
Original hinge

New hinge
Therapsid (280 mya)

Very late cynodont (195 mya)

Temporal
fenestra

Hinge Hinge
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 Figure 23.4a

Reptiles
(including
dinosaurs and birds)
OTHER
†Dimetrodon
TETRAPODS
Synapsids

Therapsids
Cynodonts †Very late (non-
mammalian)
cynodonts
Mammals

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 Synapsids (300 mya) had single-pointed teeth,
large temporal fenestra, and a jaw hinge between
the articular and quadrate bones

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 Therapsids (280 mya) had large dentary bones,
long faces, and specialized teeth, including large
canines

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 Figure 23.4b

Synapsid (300 mya) Key to skull bones

Articular
Quadrate
Temporal Dentary
fenestra Squamosal

Hinge

Therapsid (280 mya)

Temporal
fenestra

Hinge

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 Early cynodonts (260 mya) had large dentary
bones in the lower jaw, large temporal fenestra in
front of the jaw hinge, and teeth with several cusps

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 Later cynodonts (220 mya) had teeth with complex
cusp patterns and an additional jaw hinge between
the dentary and squamosal bones

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 Very late cynodonts (195 mya) lost the original
articular-quadrate jaw hinge
 The articular and quadrate bones formed inner ear
bones that functioned in transmitting sound
 In mammals, these bones became the hammer
(malleus) and anvil (incus) bones of the ear

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 Figure 23.4c
Early cynodont (260 mya) Key to skull bones
Articular
Temporal Quadrate
fenestra
(partial view) Dentary
Squamosal
Hinge

Later cynodont (220 mya)

Original hinge

New hinge

Very late cynodont (195 mya)

Hinge
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Concept 23.2: The rise and fall of groups of
organisms reflect differences in speciation and
extinction rates
 The history of life on Earth has seen the rise and fall
of many groups of organisms
 The rise and fall of groups depend on speciation
and extinction rates within the group

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 Figure 23.5

Lineage A
†
†
†
†

Common
ancestor of
lineages A Lineage B
and B
†

4 3 2 1 0
Millions of years ago
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Plate Tectonics

 At three points in time, the landmasses of Earth

have formed a supercontinent: 1.1 billion, 600
million, and 250 million years ago
 According to the theory of plate tectonics, Earth’s
crust is composed of plates floating on Earth’s
mantle

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 Figure 23.6

Crust

Mantle

Outer
core

Inner
core

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 Tectonic plates move slowly through the process of
continental drift
 Oceanic and continental plates can separate, slide
past each other, or collide
 Interactions between plates cause the formation of
mountains and islands and earthquakes

Video: Lava Flow

Video: Volcanic Eruption

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 Figure 23.7

North
Eurasian Plate
American
Juan Plate
de Fuca Caribbean Philippine
Plate Plate Plate
Arabian
Plate Indian
Cocos Plate Plate
Pacific South
Plate American
Nazca Plate African Australian
Plate
Plate Plate
Scotia
Antarctic
Plate
Plate

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Consequences of Continental Drift

 Formation of the supercontinent Pangaea about

250 million years ago had many effects
 A deepening of ocean basins
 A reduction in shallow water habitat
 A colder and drier climate inland

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 Figure 23.8
Present

Collision of
45 mya India with
Eurasia

Cenozoic
r i ca
Ame Eurasia
th
No
r
Africa Present-day
65.5 mya South India continents
America Madagascar alia
u str
A
Antarctica

Laurasia
Laurasia and
135 mya Gondwana
Mesozoic

Gon
dwa landmasses
na

a ea The supercontinent
251 mya Pa
ng
Pangaea
Paleozoic

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 Figure 23.8a

Laurasia
Laurasia and
135 mya Gondwana

Mesozoic
Gon
dw landmasses
ana

ga
e a The supercontinent
251 mya an
P Pangaea
Paleozoic

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 Figure 23.8b

Present

Collision of
45 mya India with
Eurasia
Cenozoic

a
eri c
m Eurasia
rt hA
N o Africa Present-day
65.5 mya South India continents
America Madagascar tr alia
s
Au
Antarctica

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 Continental drift can cause a continent’s climate to
change as it moves north or south
 Separation of landmasses can lead to allopatric
speciation
 For example, frog species in the subfamilies
Mantellinae and Rhacophorinae began to diverge
when Madagascar separated from India

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 Figure 23.9

Mantellinae
(Madagascar only):
100 species
Rhacophorinae
(India/southeast
Asia): 310 species

80 60 40 20 0
1 2 Millions of years ago (mya)

1 2
India

Madagascar
88 mya 56 mya

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 The distribution of fossils and living groups reflects
the historic movement of continents
 For example, the similarity of fossils in parts of South
America and Africa is consistent with the idea that
these continents were formerly attached

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Mass Extinctions

 The fossil record shows that most species that have

ever lived are now extinct
 Extinction can be caused by changes to a species’
environment
 At times, the rate of extinction has increased
dramatically and caused a mass extinction
 Mass extinction is the result of disruptive global
environmental changes

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The “Big Five” Mass Extinction Events

 In each of the five mass extinction events, more

than 50% of Earth’s species became extinct

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 Figure 23.10

1,100
25 1,000
900
(families per million years):

20 800
Total extinction rate

Number of families:
700
15 600
500
10 400
300
5 200
100
0 0
Era Paleozoic Mesozoic Cenozoic
Period E O S D C P Tr J C P N Q
542 488 444 416 359 299 251 200 145 65.5 0

Time (mya)
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 The Permian extinction defines the boundary
between the Paleozoic and Mesozoic eras 251
million years ago
 This mass extinction occurred in less than 500,000
years and caused the extinction of about 96% of
marine animal species

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 A number of factors might have contributed to these
extinctions
 Intense volcanism in what is now Siberia
 Global warming resulting from the emission of large
amounts of CO2 from the volcanoes
 Reduced temperature gradient from equator to poles
 Oceanic anoxia from reduced mixing of ocean waters

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 The Cretaceous mass extinction 65.5 million years
ago separates the Mesozoic from the Cenozoic
 Organisms that went extinct include about half of all
marine species and many terrestrial plants and
animals, including most dinosaurs

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 The presence of iridium in sedimentary rocks
suggests a meteorite impact about 65 million
years ago
 Dust clouds caused by the impact would have
blocked sunlight and disturbed global climate
 The Chicxulub crater off the coast of Mexico is
evidence of a meteorite collision that dates to the
same time

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 Figure 23.11

NORTH
AMERICA
Chicxulub
Yucatán crater
Peninsula

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Is a Sixth Mass Extinction Under Way?

 Scientists estimate that the current rate of extinction

is 100 to 1,000 times the typical background rate
 Extinction rates tend to increase when global
temperatures increase
 Data suggest that a sixth, human-caused mass
extinction is likely to occur unless dramatic action
is taken

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 Figure 23.12

3 Mass extinctions
Relative extinction rate of

2
marine animal genera

−1

−2
−3 −2 −1 0 1 2 3 4
Cooler Warmer
Relative temperature
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Consequences of Mass Extinctions

 Mass extinction can alter ecological communities and

the niches available to organisms
 It can take 5–100 million years for diversity to recover
following a mass extinction
 The type of organisms residing in a community can
change with mass extinction
 For example, the percentage of marine predators
increased after the Permian and Cretaceous mass
extinctions
 Mass extinction can pave the way for adaptive
radiations
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 Figure 23.13

50 Cretaceous
mass extinction
Predator genera (%)

40 Permian mass
extinction
30

20

10

0
Era Paleozoic Mesozoic Cenozoic
Period E O S D C P Tr J C P N
542 488 444 416 359 299 251 200 145 65.5 Q 0

Time (mya)

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Adaptive Radiations

 Adaptive radiation is the evolution of many

diversely adapted species from a common ancestor
 Adaptive radiations may follow
 Mass extinctions
 The evolution of novel characteristics
 The colonization of new regions

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Worldwide Adaptive Radiations

 Mammals underwent an adaptive radiation after the

extinction of terrestrial dinosaurs
 The disappearance of dinosaurs (except birds)
allowed for the expansion of mammals in diversity
and size
 Other notable radiations include photosynthetic
prokaryotes, large predators in the Cambrian, land
plants, insects, and tetrapods

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 Figure 23.14

Ancestral
mammal Monotremes
(5 species)

ANCESTRAL Marsupials
CYNODONT (324
species)

Eutherians
(5,010
species)

250 200 150 100 50 0

Time (millions of years ago)

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Regional Adaptive Radiations

 Adaptive radiations can occur when organisms

colonize new environments with little competition
 The Hawaiian Islands are one of the world’s great
showcases of adaptive radiation

Animation: Allometric Growth

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 Figure 23.15

Close North American

relative, the tarweed
Carlquistia muirii
KAUAI
Dubautia laxa 5.1
million MOLOKAI 1.3 Argyroxiphium
years million
OAHU
years sandwicense
3.7
MAUI
million LANAI
years
HAWAII
N
0.4
million
years

Dubautia waialealae

Dubautia scabra
Dubautia linearis
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 Figure 23.15a

KAUAI
5.1
million
years MOLOKAI
1.3 million
OAHU
years
3.7
MAUI
million LANAI
years
HAWAII
N 0.4
million
years

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 Figure 23.15b

Close North American

relative, the tarweed
Carlquistia muirii

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 Figure 23.15c

Dubautia waialealae

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 Figure 23.15d

Dubautia laxa

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 Figure 23.15e

Dubautia scabra

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 Figure 23.15f

Argyroxiphium
sandwicense

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 Figure 23.15g

Dubautia linearis

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Concept 23.3: Major changes in body form can
result from changes in the sequences and
regulation of developmental genes
 Studying genetic mechanisms of change can
provide insight into large-scale evolutionary change

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Effects of Development Genes

 Genes that program development influence the

rate, timing, and spatial pattern of changes in an
organism’s form as it develops into an adult

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Changes in Rate and Timing

 Heterochrony is an evolutionary change in the rate

or timing of developmental events
 It can have a significant impact on body shape
 The contrasting shapes of human and chimpanzee
skulls are the result of small changes in relative
growth rates

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 Figure 23.16

Chimpanzee infant Chimpanzee adult

Chimpanzee fetus Chimpanzee adult

Human fetus Human adult

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 Figure 23.16a

Chimpanzee infant Chimpanzee adult

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 Another example of heterochrony can be seen in
the skeletal structure of bat wings, which resulted
from increased growth rates of the finger bones

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 Figure 23.17

Hand and
finger bones

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 Heterochrony can alter the timing of reproductive
development relative to the development of
nonreproductive organs
 In paedomorphosis, the rate of reproductive
development accelerates compared with somatic
development
 The sexually mature species may retain body
features that were juvenile structures in an ancestral
species

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 Figure 23.18

Gills

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Changes in Spatial Pattern

 Substantial evolutionary change can also result from

alterations in genes that control the placement and
organization of body parts
 Homeotic genes determine such basic features as
where wings and legs will develop on a bird or how a
flower’s parts are arranged

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 Hox genes are a class of homeotic genes that
provide positional information during animal
development
 If Hox genes are expressed in the wrong location,
body parts can be produced in the wrong location
 For example, in crustaceans, a swimming
appendage can be produced instead of a feeding
appendage

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The Evolution of Development

 Adaptive evolution of both new and existing genes

may have played a key role in shaping the diversity
of life
 Developmental genes may have been particularly
important in this process

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Changes in Gene Sequence

 New morphological forms likely come from gene

duplication events that produce new developmental
genes
 A possible mechanism for the evolution of six-legged
insects from a many-legged crustacean ancestor
has been demonstrated in lab experiments
 Specific changes in the Ubx gene have been
identified that can “turn off” leg development

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 Figure 23.19

Hox gene 6 Hox gene 7 Hox gene 8

Ubx

About 400 mya

Drosophila Artemia

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Changes in Gene Regulation

 Changes in morphology likely result from changes

in the regulation of developmental genes rather than
changes in the sequence of developmental genes
 For example, threespine sticklebacks in lakes have
fewer spines than their marine relatives
 The gene sequence remains the same, but the
regulation of gene expression is different in the two
groups of fish

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 Figure 23.UN03

Ventral spines

Threespine stickleback
(Gasterosteus aculeatus)

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 Figure 23.20

esis A: Differences in Result: No

sequence The 283 amino acids of the Pitx1
protein are identical.
esis B: Differences in Result: Yes
expression
stickleback embryo: Lake stickleback embryo:
sion in ventral spine and expression only in mouth
regions regions

Red arrows indicate regions of Pitx1

expression.
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 Figure 23.20a

Marine stickleback embryo:

expression in ventral spine and
mouth regions

Red arrows indicate regions of Pitx1

expression.

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 Figure 23.20b

Lake stickleback embryo:

expression only in mouth
regions

Red arrows indicate regions of Pitx1

expression.

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Concept 23.4: Evolution is not goal oriented

 Evolution is like tinkering—it is a process in which

new forms arise by the slight modification of
existing forms

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Evolutionary Novelties

 Most novel biological structures evolve in many

stages from previously existing structures
 Complex eyes have evolved from simple
photosensitive cells independently many times
 Exaptations are structures that evolve in one context
but become co-opted for a different function
 Natural selection can only improve a structure in the
context of its current utility

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 Figure 23.21

pigmented cells (b) Eyecup

cells
ptors) Pigmented
cells
Epithelium

Nerve Nerve fibers

fibers
Example: Pleurotomaria, a
atella, a limpet slit shell mollusc

amera-type eye (d) Eye with primitive lens (e) Complex camera lens-
type eye
Cellular Cornea
Fluid-filled mass Cornea
cavity (lens)
Lens

Retina
Optic nerve Optic
Pigmented nerve
layer (retina) Example: Murex, a marine
autilus snail Example: Loligo, a squid

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 Figure 23.21a

(a) Patch of pigmented cells

Pigmented cells
(photoreceptors)
Epithelium

Nerve
fibers
Example: Patella, a limpet

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 Figure 23.21b

(b) Eyecup

Pigmented
cells

Nerve fibers

Example: Pleurotomaria, a
slit shell mollusc

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 Figure 23.21c

(c) Pinhole camera-type eye

Epithelium
Fluid-filled
cavity

Optic
nerve
Pigmented
layer (retina)
Example: Nautilus

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 Figure 23.21d

(d) Eye with primitive lens

Cellular Cornea
mass
(lens)

Optic nerve

Example: Murex, a marine

snail

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 Figure 23.21e

(e) Complex camera lens-

type eye

Cornea

Lens

Retina
Optic
nerve

Example: Loligo, a squid

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Evolutionary Trends

 Extracting a single evolutionary progression from

the fossil record can be misleading
 Apparent trends should be examined in a broader
context
 The species selection model suggests that
differential speciation success may determine
evolutionary trends
 Evolutionary trends do not imply an intrinsic drive
toward a particular phenotype

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 ocene

ocene
ocene
Figure 23.22

Anchitherium
Millions of years ago

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ene Oligocene Miocene

laeotherium

achynolophus

Palaeotherium
5
0

30

35

55
50
45
40
25
10

15
Equus

Sinohippus
Haplohippus
Megahippus

Hyracotherium
Mesohippus
Hypohippus
Miohippus
Pliohippus

20Merychippus

Archaeohippus

Parahippus

Hyracotherium
relatives
ihippus
Key
Grazers
Browsers

Hipparion
Neohipparion
Nannippus
Hippidion and
close relatives
Callippus
 Figure 23.22a

25 Grazers
Browsers
30

35
Mesohippus
Oligocene

Miohippus
40

Haplohippus
Palaeotherium
Millions of years ago

Pachynolophus

45
Propalaeotherium

Epihippus
50
Eocene

Hyracotherium
relatives
Orohippus

55 Hyracotherium

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 e
Millions of years ago
Figure 23.22b

Anchitherium
Miocene

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0

10

20
15
Sinohippus

Megahippus
Equus

Hypohippus
Mio-
hippus
Archaeohippus
Pliohippus

Merychippus

arahippus
Grazers
Browsers

Hipparion
Neohipparion
 Figure 23.UN02

Species with
planktonic larvae

Species with
nonplanktonic
larvae

Paleocene Eocene
65 60 55 50 45 40 35
Millions of years ago (mya)
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 Figure 23.UN04

Flies and
fleas

Caddisflies

Moths and
Herbivory butterflies

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