What is the unit of natural selection?

Individual Population Species

Evolution on the smallest scale…

Microevolution Change in allele frequencies in a

population over generations
(e.g., methicillin-resistance of hospital staph bacteria)

Change that occurs at the level of

Macroevolution
the species or higher
(e.g., the compounded effects of microevolution)

What is a species?
 The term “species” defined
Morphological Species Concept
A group of organisms with a set of prescribed similarities in
body shape and other structural features.
Useful, but Works for asexual Biological
subjective. species. basis?

Ecological Species Concept

A set of organisms exploiting a single niche
Works for Highly subjective
Operational?
asexual species (niche?)

Phylogenetic Species Concept

Smallest group of organisms that share a common ancestor
Operational? Subjective?
Good fit with (extensive (smallest
theory research) group)
 The term “species” defined (cont.)

Biological Species Concept

Groups of interbreeding natural populations that are
reproductively isolated from other such groups

Focuses on how speciation occurs

Mandates the production of fertile, viable offspring only
within a species.
Emphasizes the separateness of species based on
reproductive isolation.

Operational?
Each definition has limitations

But the BSC serves as the foundation to further explore …

 The Mechanisms of Speciation
(genetic variation, reproductive isolation, selection)
Genetic Variation
Multiple sources (for example…)

 mutations  chromosome rearrangements

 gene duplications  sexual reproduction
and…
 genetic drift

The unpredictable fluctuation of

allele frequencies in a population
from generation to generation.
 The Mechanisms of Speciation
Genetic Variation (cont.)
Let’s take a closer look at Genetic Drift…
Don’t take notes, just listen.
Flip a coin 1,000 times. An outcome of 700 heads and 300
tails would make you suspicious.

But if you flip a coin 10 times, an outcome of 7 heads

and 3 tails would not be surprising.

The smaller the number of coin flips, the more likely it

is that chance alone will cause a deviation from the
predicted result.

This is the way it is with genetic drift

and population size.
The Mechanisms of Speciation
Genetic Variation (cont.)

Consider this example from your textbook

Genetic Drift in a small population of wildflowers

Although population genetics is beyond the scope of this

course, we can summarize the importance of genetic drift…
 The Mechanisms of Speciation Genetic Variation (cont.)

4 Major Influences of Genetic Drift on Populations

Genetic drift can cause allele frequencies to change at random
Genetic drift can lead to a loss of genetic variation within
populations.
Genetic drift can cause harmful alleles to become fixed.
Genetic drift has a major impact on small populations

About 12,000 years ago, a mass

extinction event eliminated ~75% of the
world’s large mammal species. A handful
of cheetahs survived to repopulate, but
with severely reduced genetic diversity.
 The Mechanisms of Speciation Genetic Variation (cont.)

The story of the cheetah illustrates...

A sharp reduction in the size of a

Bottleneck population due to environmental
Effect events or human activities
Loss of genetic variation that occurs
Founder when a new population is established
Effect by a very small number of individuals
The Mechanisms of Speciation (cont.)

Bottleneck and founder effects…

can drastically alter allelic frequencies
can lead to the accumulation of deleterious genes
can result in the extinctions of species

~ 55 Java rhinos
remain

~ 50 Northern
Pacific right whales
remain

Directed Reading – Case Study: Impact of Genetic Drift on

the Greater Prairie Chicken
 Toba Catastrophe Theory
Highly controversial
Massive volcanic eruption ~75,000 years ago
Linked to human genetic bottleneck due to…
Sharp population decline to only 3,000–10,000
survivors
Correlated with “sudden” increase in modern
human intelligence and extinction of other human
lineages?

Lake Toba, Sumatra, Indonesia - Landsat satellite photo

 The Mechanisms of Speciation (cont.)

Reproductive Isolation
Biological factors that impede the production of fertile, viable
offspring
Blocks gene flow between “species”

Limits the formation of hybrids

Two Categories

Prezygotic
Barriers
Postzygotic
Barriers
The Mechanisms of Speciation (cont.)
Reproductive Isolation Prezygotic Block fertilization
Barriers from occurring
Act in one of three ways…
Directed Study
 Impede mating attempts Exploring
 Prevent completion of mating Reproductive
 Hinder fertilization Barriers (Ch. 24)

Five prezygotic barriers

1. Habitat Isolation
Two garter snake
species occupy
different habitats in the
same region
The Mechanisms of Speciation (cont.)
Reproductive Isolation Prezygotic
Barriers
Five prezygotic barriers (cont.)
2. Temporal Isolation

Two spotted skunk

species breed in
different seasons
The Mechanisms of Speciation (cont.) Prezygotic
Reproductive Isolation
Barriers
Five prezygotic barriers (cont.)
3. Behavioral Isolation Unique behavioral
rituals may be
required for mate
recognition.

Blue-footed booby
The Mechanisms of Speciation (cont.) Prezygotic
Reproductive Isolation
Barriers
Five prezygotic barriers (cont.)
4. Mechanical Isolation Morphological
differences prevent
successful mating
The Mechanisms of Speciation (cont.) Prezygotic
Reproductive Isolation
Barriers
Five prezygotic barriers (cont.)
5. Gamete Isolation Sperm and eggs may
not be compatible
The Mechanisms of Speciation (cont.)
Postzygotic
Reproductive Isolation
Barriers
Occur after fertilization when prezygotic barriers fail
Three postzygotic barriers… Interspecies offspring
1. Reduced hybrid viability may have incomplete
development or
reduced vigor

Though subspecies of Ensatina

have overlapping ranges, they
cannot produce viable hybrids.
The Mechanisms of Speciation (cont.)
Postzygotic
Reproductive Isolation
Barriers
Three postzygotic barriers (cont.)
2. Reduced hybrid fertility Hybrids may be
Mule vigorous, but sterile.
Zorse

Liger
The Mechanisms of Speciation (cont.)
Postzygotic
Reproductive Isolation
Barriers
Three postzygotic barriers (cont.)
3. Hybrid breakdown Hybrids are fertile, but
subsequent offspring
Cotton are feeble or sterile
hybrids
rice
strains

2-spotted
spider mites
 Modes of selection
Natural selection can alter the frequency
distribution of heritable traits in three ways…

Directional
Selection
Disruptive
Selection

Stabilizing
Selection
 Modes of selection (cont.)
Directional Selection

Shift in population
traits by favoring
variants at one
extreme

Selective
pressures
against the
lighter
phenotypes
 Modes of selection (cont.)
Disruptive Selection

Shift in population
traits by favoring
variants at both
extremes

Selective
pressures
against the
intermediate
phenotypes
 Modes of selection (cont.)
Stabilizing Selection

Shift in population
traits by favoring the
intermediate
variants

Selective
pressures
against the light
and dark
phenotypes
 Directional Selection?

1850s

1960s

Has occurred twice: First in one direction and then

back in the other direction
Sexual Selection
Selection driven by the
competition for mates

Ornaments, armaments,
and behaviors
Directed Study: Sexual Selection in Gray Tree Frogs (Fig. 23.16)
 Speciation in the Presence and Absence of
Geographic Barriers

Allopatric Speciation
Speciation mediated by barrier-interrupted gene flow
Geographic speciation (synonym)
The most common method by which new species arise

Different selection
pressures

Subpopulations Two separate

become isolated species w/o barrier
Allopatry (cont.)

Speciation of antelope squirrels on

opposite rims of the Grand Canyon
 The Kaibab squirrel is a distinct
subspecies attributed to 10,000 years of
geographic isolation on the South Rim
 The subfamily of
mantella frogs is
unique to
Madagascar

How did this

happen?
Divergence began 88 mya with the separation of
Madagascar from the Indian landmass
Directed Study: Allopatry and reproductive
isolation in fruit flies (Fig. 24.7)
Speciation in the Presence and Absence
of Geographic Barriers (cont.)

Sympatric Speciation
Speciation that occurs in geographically
overlapping populations
Less common and less
likely than allopatry
Requires mechanisms
that reduce gene flow
Mechanisms
 chromosomal changes
 nonrandom mating
Sympatric Speciation
Polyploidy in plants
Errors in cell division increases chromosome number
generating gamete isolation or reduced hybrid viability

Gametes with
Mitotic error doubles double the
chromosome number chromosome Fertilization results
number in tetraploidy

more common plants

only polyploids can interbreed
Goatsbeard (Pacific NW)
Common technique in plant
breeding
Sympatric Speciation
Habitat Differentiation

Exploitation of distinct resources (subhabitats) may select for

differences in subpopulations, affecting nonrandom mating

Emerging species of apple maggot fly

Two different habitats (hosts)
New host  increases development  earlier mating
 Directed Study: Sympatry and Sexual
Selection in Lake Victoria Cichlids (Fig. 24.12)
 The Time Course of Speciation
Two schools of thought…
evolution occurs uniformly by
Phyletic Gradualism steady, measured transformation
(syn., “anagenesis”)

Time

In recent years, anagenesis has not been well

supported by scientific evidence
The Time Course of Speciation (cont.)
Second school of thought…
Evolution exhibits long periods of
Punctuated Equilibrium stasis with rare, localized, rapid
events of branching speciation

Time

Recent evidence suggests that punctuated equilibrium

may be the general pattern of evolution

New species appear to form rapidly

Phyletic Gradualism Punctuated Equilibrium
 Let’s consider
origins and
evolutionary trees.