Learning Goal: Evaluate the relationship

between a fitness landscape and trait

evolution in a population.
Students will be able to:
• Evaluate how changes in fitness due to natural
selection change allele and genotype frequencies
• Interpret fitness curves and landscapes
• Explain how adaptive traits and non-adaptive traits
can both be present in a population
• Predict how different mechanisms of evolution may
alter a population’s allele and genotype frequencies,
and its placement on a fitness landscape
 Let’s think about how changes in an
individual’s fitness may affect allele and
genotype frequencies in a population
over time.
 Fitness Curves

Fitness curves show

us the most fit values
for a trait in a given
environment
 Fitness Curves

Fitness curves show

us the most fit values
for a trait in a given
environment

What are we really inferring from this graph?

 Fitness Curves

Fitness curves show

us the most fit values
for a trait in a given
environment

In 1978 on Daphne
Major, most ground
finches had beak depths
around 9.8 mm
 Fitness Curves
Recall the drought that
hit Daphne Major in
1977. Did it influence
mean trait values
between 1977 & 1978?
A. True
B. False

In 1978 on Daphne
Major, most ground
finches had beak depths
around 9.8 mm
 Fitness Curves

Number of individuals with

a certain trait value (or the
frequency of that value) is
observed directly & used
as a proxy for fitness

In 1978 on Daphne
Major, most ground
finches had beak depths
around 9.8 mm
 Fitness Curves
Fitness curves plotted in 2 dimensions illustrate
the fitness of a single trait…
Individuals/Fitness
Number of

Trait measurement

…but selection acts on the individual, and individuals have many

traits controlled by multiple genes and epistatic interactions.
Beak Morphology and Gene Expression

Geospiza fortis

BMP4 Expression

BMP4 Expression in
chick embryos

Normal Expression Overexpression

 Many genes contribute to an organism’s
survival and reproductive success

• Selection on a trait
doesn’t occur in a
vacuum.
• Individuals don’t pass
on just one trait to their
offspring, but rather
half their genome –
which has genes for
other traits as well.
 Many genes contribute to an organism’s
survival and reproductive success
Selection favoring one trait may constrain another
Consider body size:
- large-bodied animals
are less vulnerable to
predation
- but large bodies also
require more resources
(space, food)
 Channel Islands foxes are descended from
gray foxes of the mainland Americas

gray fox
Channel Islands fox
Urocyon cinereoargenteus
Urocyon littoralis

They look very similar to each other, but…

…Channel Islands foxes evolved smaller body sizes in
isolation from their ancestors

Which mode of selection began acting upon island fox

body size immediately after island colonization?

A. directional B. disruptive C. stabilizing

gray fox Channel Islands fox

If we assume stabilizing selection maintains a balance
between predation avoidance and energy requirements
in mainland gray foxes, why are their descendants
smaller on the Channel Islands?

A. resources are B. islands often C. both of these factors

more limited lack mainland likely contribute to
on islands predators small body size

gray fox Channel

Islands fox
Reduced body size caused by isolation is seen in
other species and is called “island dwarfism”

The pygmy mammoth also became isolated on the

Channel Islands and was much smaller than its
mainland relatives
 In contrast, smaller animals often grow larger on islands
due to decreased competition and lack of predation

The giant weta is found on islands off the coast of New Zealand
 Fitness Landscapes
We can visualize the fitness of two interacting
traits using 3-dimensional plots

Fitness

Plots can take many different forms, often with multiple

peaks & valleys – collectively called a fitness (or
adaptive) landscape
What does this wide-topped dome show us about the most fit
combination of birth weight & gestation in humans?

A. one trait combination is

much more fit than all
others
B. a range of trait
combinations have
nearly equal fitness
C. multiple trait
combinations of peak
fitness are separated
by valleys of less fit
trait combinations
 How would the appearance of this fitness landscape be
different if one precise combination of birth mass & gestation
was much more fit than all others?

A. dome would narrow to a

sharp peak
B. dome would expand to
a wider top surface
C. internal valleys would
result in multiple peaks
 Fitness Landscapes
Adaptive landscapes can often have multiple peaks

trait/gene interactions result in combinations w/ high fitness

(“peaks”) separated by combinations w/ low fitness (“valleys”)
 Fitness Landscapes
If a population’s trait values start at point X on this fitness
landscape, where do you expect it to end up after many
generations (assuming a stable environment)?

A B

D. It can move
anywhere on
X the landscape

C
 Fitness Landscapes
Fitness peaks on an adaptive landscape are often different
heights, resulting in mutiple trait combinations with high fitness

Peak
Peak 2
1

A population may become “trapped” on one peak despite

the potential for higher fitness on another peak
 Fitness Landscapes
Discuss with a neighbor: How might a population “jump”
across a valley from one fitness peak to another higher one?

?
***
*
If a population takes the path shown on the adaptive
landscape below, which mode of selection does this
best represent?

A. stabilizing
B. directional
C. disruptive
What might you expect if a population climbed both
peaks on this adaptive landscape simultaneously?
A. population that climbs the lower peak will go extinct
B. population that climbs the lower peak will “jump” to
the higher peak
C. genetic divergence would ultimately result in two new
species
 For speciation to occur, two populations must be
reproductively isolated. What would you predict if
gene flow continued between all individuals on this
fitness landscape?
Next time – Speciation!!!
Speciation
Learning Goal: Recognize species formation as the result of
evolutionary mechanisms and that species interactions can contribute
to this process.
Upon completion of this material, students should be able to:
• Differentiate between the various species’ concepts and the pros and cons
of applying them.
• Describe different models of speciation, with examples.
• Define a population and how a population is distinct from a species.
(Discuss the evolutionary significance of this distinction.)
• Describe the term fitness and its impact on evolutionary processes.
• Apply a molecular clock to estimate time since divergence of two species.
• Interpret data from a fitness landscape and how this may affect
evolutionary trajectories in a given species, and lead to speciation
• Recognize that adaptation is not the result of organismal need
• Determine the role of drift, natural selection, and mutation in divergence
events leading to speciation
• Relate the pre- and post-zygotic isolating mechanisms to the process of
speciation
 Life is Squishy
• Speciation can be a complicated process that often takes time

• For this reason, species themselves can sometimes be difficult to

distinguish

• Species concepts are created by us, and no one species concept fits
all situations

• The impossibility of using a single set of criteria to define all species

is one thing that illustrates how evolution can work in many different
ways to produce many different forms and functions

• Despite this difficulty, there are several lines of evidence that can be
used to identify species and several well-known mechanisms for how
the speciation CAN occur depending on the circumstances
 First you must define
“species”
•Morphological
•Paleontological
•Ecological
•Phylogenetic
•Biological
Morphological concept
of “species”
Paleontological
concept of
“species”
Ecological concept of
“species”
Phylogenetic concept of
“species”
 Biological
concept of
“species”
 Prezygotic reproductive
isolation
•Habitat isolation
•Temporal isolation
•Behavioral isolation
•Mechanical isolation
•Gametic isolation
Prezygotic reproductive
isolation - habitat
Prezygotic reproductive
isolation - temporal
Prezygotic reproductive
isolation - behavioral
Prezygotic reproductive
isolation - mechanical
Prezygotic reproductive
isolation - gametic
 Postzygotic reproductive
isolation
•Reduced hybrid viability
•Reduced hybrid fertility
•Hybrid breakdown
Postzygotic reproductive isolation –
reduced hybrid viability
Postzygotic reproductive isolation –
reduced hybrid fertility
 Postzygotic reproductive
isolation – hybrid breakdown
Imagine a population of
voles.
• The population
covers a large area of
land.

• The soil, rainfall,

amount of sunlight,
and herbivores
throughout the voles’
distribution are the
same.
The original population is split in two when a
nearby river changes course and flows
through the middle of the population
Population West and East are now
separate species.

A. true
B. false
C. need more
information
 As soon as the two populations are
separated, which of the following will
have the greatest effect on divergence
between the two populations?

A. reproductive isolation
B. genetic drift
C. natural selection
D. gene flow
Genetic differences begin to accumulate between the
two populations, and the number of unique alleles
between the populations will increase over many
generations.
 What was the origin of the unique
alleles in each of the subpopulations?
A. Gene flow from occasional immigrants from the
other subpopulation
B. Mutations that occur randomly throughout the
genomes of individuals in each subpopulation
C. Nonrandom mating within each subpopulation
D. Reduced population size in each subpopulation
after the river’s course changed
What is the significance of the accumulating
genetic differences between West and East?
A. The two populations are becoming
phylogenetically distinct from each other.
B. The two populations are reproductively
isolated from each other.
C. If the river changes course and they are
reunited, the resulting population will have
more variation and they will interbreed.
Genetic differences continue to accumulate between the
two populations, and the number of unique alleles
between the two populations continue to increase.
 Eventually, enough time has passed that
West and East are genetically distinct from
one another, and they can no longer
interbreed. Which form of speciation has
occurred?

A. allopatric speciation
B. sympatric speciation
C. peripatric speciation
What happened to
the original
population?
Talk with your classmates and draw a
picture showing what has occurred
Does your picture look
something like this?

East

West
What happened to the original
population?

East

West
What is occurring along each
of these two branches?

East

West
New species arise when two
populations have diverged
sufficiently from one another
 Reproductive isolation

Is there more than one way to achieve reproductive isolation?

Pre-zygotic and Post-zygotic isolating mechanisms
Reproductive isolation

How do you think reproductive isolation

 Back to the Voles…

A new species of grass with much

larger seeds appears on the east
side of the river. Found inside the
black circles on the map.

Some of the voles begin to only eat

larger seeds of the new grass
species while other voles continue to
feed on smaller native seeds.

During mating season, voles find

mates in the same places that they
feed.
 Which form of selection would you
expect in population East for surface
area for grinding on teeth
(for eating the new seeds)?

A. disruptive selection
B. stabilizing selection
C. directional selection
Variation in tooth morphology before
and after introduction of the new
grass species in population East
 Individuals only mate with other
individuals that eat the same size seeds
in their population. Over time, two new
species, East1 and East2, develop from
the original eastern population. This is the
result of ________ speciation.

A. allopatric
B. sympatric
C. peripatric
How does this fitness landscape relate to the
changes we see within the East population?

Peak 2
Peak 1
When individuals from species East1 and East2
mate, their offspring are very small and unable to
eat seeds of any size. As a result, the hybrid
offspring die before they can produce offspring.
This is an example of:

A. a pre-zygotic isolating mechanism.

B. a post-zygotic isolating mechanism.
C. reinforcement.
D. sexual selection.
 Reinforcement is

A. selection against hybrid offspring.

B. selection against individuals whose hybrid
offspring have reduced fitness relative to non-
hybrids.
C. pre-zygotic selection against hybrids because
they aren’t able to breed.
Reinforcement

• Post-zygotic isolation
happens first
• Reinforcement facilitates
pre-zygotic isolation
because post-zygotic
isolation causes
reinforcement to happen
• So…reinforcement is
selection for assortative
mating which avoids
hybridization
If hybrids formed between individuals from
East1 and East2 and had equal fitness to
either parental species, then this would:
A. increase the rate of speciation between East1
and East2.
B. not affect the process of speciation between
East1 and East2.
C. prohibit speciation between East1 and East2.
 Male frogs give calls that attract female frogs to
approach and mate. Researchers examined mating
calls of closely related tree frogs in South America. If
reinforcement of post-zygotic isolation is occurring,
what would you expect if you compare the calls of the
two species in zones of sympatry versus zones of
allopatry?

A. Calls would be more different in areas of sympatry.

B. Calls would be more similar in areas of sympatry.
C. Calls would be about the same in both areas.
D. There is not enough information to decide what is
happening.