Descent with Modification

Evolution
 1809
Lamarck publishes his
hypothesis of evolution.

1798 Sketch of a flying

Malthus publishes 1812 frog by Wallace
“Essay on the Principle Cuvier publishes his
of Population.” extensive studies of 1858
vertebrate fossils. While studying species in the
1795 Malay Archipelago, Wallace
Hutton proposes 1830 (shown above in 1848) sends
his principle of Lyell publishes Darwin his hypothesis of
gradualism. Principles of Geology. natural selection.

1790 1870
1809 1831–1836 1859
Charles Darwin Darwin travels On the Origin of
is born. around the world Species is published.
on HMS Beagle. 1844
Darwin writes his
essay on descent
with modification.
Marine iguana
in the
Galápagos
Islands
 Influential Folks
• Carl von Linnaeus
– Father of Taxonomy
• Organisms can be classified hierarchically
• binomial nomenclature
• James Hutton
– This geological gradualism takes long periods of time
• Thus earth is older that the 5ooo years
• Charles Lyell (took Hutton’s work)
– Took Hutton’s theory of gradualism and proposed
uniformitarianism
• the earth was shaped by
slow-moving forces still in
operation today
 Thomas Malthus
*Theorized that overproduction of populations led to limited resources
*Introduced concept of carrying capacity and natural selection

www.mnsu.edu/emuseum/biology/evolution
 Georges Cuvier
• Father of vertebrate paleontology - “Essay on the
Theory of the Earth”
– While excavating Cuvier found that the older the strata the
less the fossilsin that stratum looked like current life-forms.
– Theory of Catastrophism
• viewed the geological history of the earth as the alternation of long periods of
relative calm with comparatively brief, catastrophic events that greatly changed the
face of the planet.
 Darwin’s Observations Lead to New
Ideas
• Adaptations
– Animals gain characteristics
that enhances their survival
and reproduction in a given
environment.
• Adaptations arise through
natural selection
– Individuals with certain
inherited traits leave more
offspring than those with
other traits.
 The Origin of Species

• Descent with modification is the

explanation for life’s unity and diversity
• Natural selection is the mechanism

• Result of inferences drawn from

observations.
Darwin’s Observations of Nature

And the inferences he drew from

them
 Fig. 22-9

Terminal
Observation: bud
Artificial Selection Lateral
buds

Cabbage Brussels sprouts

Flower
clusters

Leaves

Kale
Cauliflower

Stem

Wild mustard

Flowers
and stems
Broccoli Kohlrabi
Observation
Variability in
traits within a
population
 Observation
-Traits are
inherited from
parent to child.
-No two of the
same species are
exactly alike even
if they have the
same parents
Observation
All species are
capable of
reproducing
beyond the
support of the
environment
 Observation: Many offspring do not survive due to
limitation of the environment.

• Thus, by inference, more fit individuals will leave

more offspring than less fit individuals

• Unequal ability of individuals to survive and

reproduce leads to an accumulation of favorable
traits
-- takes generations
 Natural Selection: A Summary
• Individuals with certain heritable traits survive and
reproduce at a higher rate than other individuals.
• Natural selection increases the match between
organisms and their environment over time
• If an environment changes over time, natural
selection may result in adaptation to these new
conditions and may give rise to new species.
– Note that individuals do not evolve; populations evolve
over time
• Natural selection can only increase or decrease
heritable trait that vary in a population
• Adaptations vary with different environments
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

1 A small population of birds

flies from the South American
mainland, where they fed on
seeds of a variety of sizes,
and become residents of a
distant island.

2 The birds produce many

offspring that vary in beak
size. The variation is due to
random mutations within
genes that affect beak size.

3 Due to limited resources,

not all offspring reproduce.
Surviving The seeds on this island are
birds that relatively large. Those
reproduce offspring that happen to
have larger beaks are better
at crushing these seeds, so
they are more likely to
survive and reproduce.

4 The birds of the next

generation tend to have
larger beaks.

5 After many, many generations,

the adaptation that allows
success in feeding on larger
seeds has created a new
species with larger beaks, as
well as other modified traits,
such as changes in color, that
are suited to the new
15 environment.
16
 Homology
• Homology is similarity resulting from common
ancestry and is the result of divergent
evolution.
• Homologous structures are anatomical
resemblances that represent variations on a
structural theme present in a common ancestor

Anatomical Homology
 A Different Cause of Resemblance:
Convergent Evolution
• Convergent evolution is the evolution of similar, or
analogous, features in distantly related groups
• Analogous traits arise when groups independently adapt to
similar environments in similar ways
• Examples:
 Giant anteater and echidna both have long snouts and
tongues to feed on ants
 Aerial rootlets for clinging in English ivy and
wintercreeper
 Antifreeze proteins in different, very cold water fish

• Convergent evolution does not provide information about

ancestry.
 The Molecular Processes That
Underlie Evolution

• Homologous genes– two genes derived

from the same ancestral gene
Orthologs occur in separate species

• Reveals molecular details of evolutionary

change
• Two sequences may be similar, but not
identical due to the independent
accumulation of different random mutations.
19
What is Speciation?
• Speciation, the origin of new species, is at the
focal point of evolutionary theory
• Evolutionary theory must explain how new
species originate and how populations evolve
• Microevolution consists of changes in a
single gene in a population over time
• Macroevolution refers to formation of new
species or groups of species
• Microevolution is a change in allele frequencies
in a population over generations
– Change because
Introduce new genetic variation (mutations, gene
duplication, exon shuffling, horizontal gene transfer)
– Not a major factor dictating allele frequencies
• Three mechanisms cause allele frequency change
– Natural selection
– Random genetic drift
- gene flow through migration and nonrandom mating
*only natural selection causes adaptive evolution
 Populations and Hardy Weinberg
Equillibrium
• A population is in Hardy-Weinberg equilibrium if the
genotype frequencies are the same in each generation.
• Conditions that must be met include
– There is no migration (“gene flow”) in or out of the population
– Natural selection is not occuring
– Mutation is not accruing
- Each member of the population is equally likely to breed
– The populationists infinitely large
• As long as a population satisfies biological conditions the
allele
• Frequencies (p and q)are the same in each generation. No
changes in their frequencies but we know that there can be
changes.
 Agents that Change Allele Frequency
in a Population
• Natural Selection
– Some are more successful than others in surviving and
reproducing owing to traits that give them a better fit to their
environment.
• Relative fitness is the contribution an individual makes to the gene
pool of the next generation, relative to the contributions of other
individuals
– includes intrasexual and intersexual (mate choice) sexual selection
– Causes Adaptive evolution
• Leads to greater relative fitness
– Outcomes of natural selection
• -Directional selection
• -Disruptive selection
• -Stabilizing selection
• -Balancing evolution
 Fig. 23.13

Initiators
New allele with higher fitness
introduced
Prolonged environmental change

Directional Disruptive Stabilizing

Likely to occur in populations

that occupy heterogeneous environments
 Balancing selection
• Maintains genetic diversity
• Balanced polymorphism (always subject to change
given what the environment is, but takes long time
for the process of change to occur)
Two or more alleles are kept in balance, and therefore
are maintained in a population over
the course of many generations
• Two common ways
For a single gene, heterozygote favored
• Heterozygote advantage – HS allele
Negative frequency-dependent selection
• Rare individuals have a higher fitness
26
Why Natural Selection Cannot Fashion
Perfect Organisms
1. Selection can act only on existing variations
(act on existing genes)
2. Evolution is limited by historical constraints
(ex; big bang theory)
3. Adaptations are often compromises
(adapting to given environment because
there is always changes in specific alleles)
4. Chance, natural selection, and the
environment interact
 Natural selection is the only
mechanism that consistently causes
adaptive evolution
• Evolution by natural selection involves both chance
and “sorting”
– New genetic variations arise by chance
– Beneficial alleles are “sorted” and favored by natural
selection
• Only natural selection consistently increases the
frequencies of alleles that provide reproductive
advantage
• Natural selection brings about adaptive evolution
by acting on an organism’s phenotype.
 Genetic drift = unpredictable changes in
allele frequency in a small population
Genetic drift is significant (and faster) in small
populations
1. Causes allele frequencies to change at random
1. Unrelated to fitness
2. Genetic drift can lead to a loss or fixation of an
allele (Frequency = 0 or 100%)
3. Genetic drift can cause harmful alleles to
become fixed
 Agents that Change Allele Frequency in a Population
• Genetic drift
– Founder Effect
• Small number of individuals become isolated from larger population
– Establishes different gene pool from population due to limited variation

– Bottleneck
– Disaster strikes. Chance alone leads to certain alleles being more or
less present in survivors
– Migration and Nonrandom mating
• One member of population is not equally likely to mate with any other
member.
– can reduce genetic variation in populations, potentially reducing its
ability to evolve in response to new selective pressures.
– can have drastic results in smaller populations.
» Endangered species.
– can contribute to speciation.
» a small isolated population may diverge from the larger population
 Nonrandom mating
• Forms of nonrandom mating
Assortative
Individuals with similar phenotypes are more likely to mate
Increases the proportion of homozygotes

Disassortative mating
Dissimilar phenotypes mate preferentially
Favors heterozygosity
Inbreeding
Does not favor any particular allele but does increase the
likelihood the individual will be homozygous
May have negative consequences with regard to recessive
alleles
31
 Agents that Change Allele Frequency
in a Population
• Gene flow consists of the movement of alleles
among populations
– Immigration
– Emigration
• Alleles can be transferred through the
movement of fertile individuals or gametes (for
example, pollen)
– Gene flow tends to reduce differences between
populations over time
• Gene flow is more likely than mutation to alter
allele frequencies directly
 Agents that Change Allele Frequency
in a Population
• Mutation
– A single mutation can have a large effect, but in many
cases, evolutionary change is based on the
accumulation of many mutations.
– must occur in germ line
• Sex
– introduces new gene combinations into a population.
• type of genetic shuffling that is a source of genetic
variation.
Macroevolution

What is a species?
 Morphological traits
• Physical characteristics of an organism
• Drawbacks for determining species
How many traits to consider
Traits may vary in a continuous way
What degree of dissimilarity to use
Members of the same species can look very
different
 Members of a different species can look very
similar

35
 The Biological Species Concept
• The biological species concept states that a species is a
group of populations whose menbers have the
potential to interbreed in nature and produce viable,
fertile offspring; they do not breed successfully with
other populations
• Gene flow between populations holds the phenotype
of a population together
 Reproductive isolation
Prevents one species from successfully
interbreeding with other species
Four main problems for determining species
– May be difficult to determine in nature
– Can interbreed and yet do not
– Does not apply to asexual species (particularly in
unicellular species)
– Cannot be applied to extinct species.

37
 Molecular features
• Compare features to identify similarities
and differences among different
populations
DNA sequences within genes
Gene order along chromosomes
Chromosome structure
Chromosome number
• May be difficult to draw the line when
separating groups

38
 Ecological factors
• Variety of factors related to an organism’s habitat
can be used to distinguish one species from
another
• Many bacterial species have been categorized as
distinct species based on ecological factors
Drawback – different groups of bacteria sometimes
display very similar growth characteristics, and even
the same species may show great variation in the
growth conditions it will tolerate

39
• Evolutionary lineage concept
Species should be defined based on the
separate evolution of lineages

• Ecological species concept

Each species occupies an ecological niche–
the unique set of habitat resources that a
species requires, as well as its influence on the
environment and other species

40
 Species concepts

 Way to define the concept of a species and/or

provide an approach to distinguish one species
from another
 Biological species concept
– Species is a group of individuals whose members have
the potential to interbreed with one another in nature
to produce viable, fertile offspring
– But cannot successfully interbreed with members of
other species

41
 Biological Species Concept
Reproductive isolation is the existence of biological factors (barriers) that impede two species
from producing viable, fertile offspring

Reproductive isolation can be classified by whether factors act before or after fertilization

Hyprids are the offspring of crosses between different species

Prezygotic barriers Postzygotic barriers
Habitat Temporal Behavioral Mechanical Gametic Reduced Hybrid Reduced Hybrid Hybrid
Isolation Isolation Isolation Isolation Isolation Viability Fertility Breakdown

Individuals VIABLE,
of MATING
different FERTILIZATION FERTILE
ATTEMPT
species OFFSPRING

(a) (c) (e) (f) (g) (h) (i) (l)

(d) (j)

(b)

(k)
 Limitations of the Biological Species
Concept
• The biological species concept cannot be applied
to fossils or asexual organisms (including all
prokaryotes).
• The biological species concept emphasizes
absence of gene flow
• However, gene flow can occur between distinct
species
– For example, grizzly bears and polar bears can
mate to produce “grolar bears”
 Cladogenic Speciation
• occurs when a population is different enough
from its ancestral species so that no genetic
exchange can occur between them
– the appearance of a reproductive isolating
mechanism produces 2 or more species where
there was one
• Two modes
– Allopatric speciation
– Sympatric speciation
 Allopatric Speciation
• Speciation that takes place in populations
with geographically separate
ranges. Gene flow is interrupted and new
species evolve.
• The separated population encounters
different selective pressures and its gene pool
is subjected to bottleneck
 Sympatric speciation
• Occurs when members of a species that
are within the same range diverge into
two or more different species even though
there are no physical barriers to
interbreeding
• Mechanisms include
Polyploidy
Adaptation to local environments
Sexual selection (mating behaviors)
46
 Adaptation to local environments
– Geographic area may have variation so that some
members of a population may diverge and occupy
different local environments that are continuous with
each other (organisms adapting to preferred
area/environment)

 Sexual selection
– Certain females prefer males with one color pattern,
while other females prefer males with a different color
pattern (over a long period of time can lead to different
species).

47
 Sympatric Speciation
• Disruptive selection
– Animals diverge mostly due to reproductive
isolation.
– Reproductive isolation is a result of genetic
factors that cause offspring to rely upon
resources not used by previous
generations. (Example: switch to a new food
source
• Example: Lake Victoria has 200 closely related species
of Cichlids (fish) which probably all arose from one
ancestor
 Hybrid zones reveal factors that cause
reproductive isolation
• A hybrid zone is a region in which members of
different species mate and produce hybrids
• Hybrids are the result of mating between
species with incomplete reproductive barriers
 Reinforcement Fusion and Stability
• When hybrids are less fit than parent species,
reinforcement of reproductive barriers may occur
through strong selection for prezygotic barriers
– Over time, the rate of hybridization decreases
• Fusion: If hybrids are as fit as parents, there can be
substantial gene flow between species
– If gene flow is great enough, reproductive barriers
weaken and the parent species can fuse into a single
species
• Stability: Extensive gene flow from outside the
hybrid zone can overwhelm selection for increased
reproductive isolation inside the hybrid zone
 Adaptive Radiation
• produced from a combination of allopatric
and sympatric speciation
– separation from ancestral species allopatrically
– presence of a variety of open ecological niches
provides opportunity for sympatric speciation by
disruptive selection
 Adaptive Radiations
• Adaptive radiation is the rapid evolution of
diversely adapted species from a common
ancestor
• Adaptive radiations may follow
– Mass extinctions
– The evolution of novel characteristics
– The colonization of new regions
• Adaptive radiations can occur when organisms
colonize new environments with little
competition.
 The Pace of Speciation
• Gradualism
 Each new species evolves continuously over long
spans of time
 Large phenotypic differences that produce new species
are due to the accumulation of many small genetic
changes

• Punctuated equilibrium
 Tempo more sporadic
 Species in equilibrium for long periods and then
short rapid bursts of changes

• Both views have merit

53
 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 (fins to
legs)
– Natural selection can only improve a structure in the
context of its current utility (does not invent but
improves)
 The CHALLENGE

How do we organize the tremendous

diversity of living organisms?
 Taxonomy

• Taxonomy
Science of describing, naming, and
classifying living and extinct organisms and
viruses
• Systematics
Study of biological diversity and the
evolutionary relationships among organisms,
both extinct and modern
* Systematists use fossil, molecular, and
genetic data to infer evolutionary relationships.
56
 Taxonomy
• Hierarchical system involving successive
levels

• Each group at any level is called a taxon

• Highest level is Domain

All of life belongs to one of 3 domains
Bacteria, Archaea, and Eukarya

57
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Taxonomic Gray wolf Number of

group found in species

Domain Eukarya ~4 –10 million

Supergroup Opisthokonta >1 million

Kingdom Animalia >1 million

Phylum Chordata ~50,000

Class Mammalia ~5,000

Order Carnivora ~270

Family Canidae 34

Genus Canis 7

Species lupus 1

58
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Domains: Bacteria Archaea Eukarya

Eukaryotic
supergroups:
Excavata Land plants and relatives Alveolata Stramenopila Rhizaria Amoebozoa Opisthokonta

Typical protists:

Large eukaryotic Plantae Fungi Animalia

kingdoms:

59
 Linnaeus convinced Darwin provided us with
us to use a the mechanism by which
hierarchical evolution results in
classification descent with modification
system

Taxonomy – naming & classifying organisms

Systematics – naming & classifying
organisms according to their evolutionary
relationships -Systematic
Phylogenetics – reconstructing the Phylogenetics
evolutionary relationships among
organisms
 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Species F, I, G, J, H, and K at the tips of branches in the present are extant species that still exist.

Species B and E at the

tips of branches in the
F I G past are extinct species. J H K
Present

Anagenesis:
Species C
evolved into
species G. B E
Millions of years ago (mya)

5
C
D
Clade: This group
Time

includes all of the

species that were
derived from the
ancestor, species D.
B
10
Cladogenesis:
Species A diverged
A into species A and B.
Branch points or nodes indicate
when a species diverged into 2
or more different species.

61
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H I J K L M N O H I J K L M N O H I J K L M N O

D E F G D E F G D E F G

B C B C B C

A A A
A monophyletic A paraphyletic A polyphyletic
group contains a group contains a group contains
common ancestor common ancestor groups of species
and all of its but not all of its with different
descendants. descendants. common ancestors.

(a) Monophyletic group (b) Paraphyletic group (c) Polyphyletic group

62
 Cladistic approach compares homologous traits, also called
characters, which may exist in two or more character states
 Shared primitive character or symplesio-morphy
– Shared by two or more different taxa and inherited from
ancestors older than their last common ancestor
 Shared derived character or synapo-morphy
– Shared by two or more species or taxa and has originated in
their most recent common ancestor
– Basis of the cladistic approach is to analyze many shared
derived characters to deduce the pathway that gave rise to
those species

63
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With regard to species D and E, having 2 eyes is a shared

primitive character, whereas having 2 front flippers is a
shared derived character.

D E F G

B C

2 eyes,
2 eyes, 2 front legs
2 front flippers

2 eyes,
A
2 front legs

64
 Constructing a cladogram

1. Choose species

2. Choose characters

3. Determine polarity of character states

 Primitive or derived?

65
4. Analyze cladogram
 All species (or higher taxa) are placed on tips
in the phylogenetic tree, not at branch points
 Each cladogram branch point should have a
list of one or more shared derived
characters that are common to all species
above the branch point unless the character
 All shared derived characters appear
together only once in a cladogram unless
they arose independently during evolution
more than once
 Principle of parsimony
Preferred hypothesis is the one that is the
simplest for all the characters and their states
Challenge in a cladistic approach is to
determine the correct polarity of states
It may not always be obvious which traits are
primitive (came earlier) and which are derived
(came later in evolution)
– Fossils may be analyzed to help resolve

67
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Lancelet Lamprey Salmon Lizard Rabbit

Mammary
glands

Tetrapod
Lancelet Lamprey Salmon Lizard Rabbit

Notochord Yes Yes Yes Yes Yes

Hinged jaw
Vertebrae No Yes Yes Yes Yes
Hinged jaw No No Yes Yes Yes
Tetrapod No No No Yes Yes Vertebrae
Mammary No No No No Yes
glands
Notochord
(a) Characteristics among species
(b) Cladogram based on morphological traits

68