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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014 Lecture 3 Lecture 3: Characters, Traits and States BIS 002C Biodiversity & the Tree of Life Spring 2014 Prof. Jonathan Eisen 1

BIS2C. Biodiversity and the Tree of Life. 2014. L3. Characters, Traits and States

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3

!

Lecture 3: !

Characters, Traits and States !!!

BIS 002C Biodiversity & the Tree of Life

Spring 2014 !

Prof. Jonathan Eisen

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Where we are going and where we have been

• Previous Lecture: !Trees, Taxa and Groups.

• Current Lecture: !Characters, Traits and States

• Next Lecture: ! Inferring trees.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 2 Outline

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• Groupings in trees !Monophyletic groups !Non monophyletic groups !Relatedness ! Ingroups and outgroups ! “Sisters”

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

Sister groups: two clades or species that are each other’s closest relatives (the two descendants of a single node)

Key Concept: Sister groups

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

g and h are “sisters” or “sister taxa”

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Key Concept: Sister groups

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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The group “g and h” is sister to f

f is sister to the group “g and h”6

Key Concept: Sister groups

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3: Key Concept

Using a phylogenetic tree to trace and infer the history of traits.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3: Outline

• Knowing history

• Character traits and states

• State changes

• Synapomorphies

• Homoplasy

• Character state reconstruction

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3: Outline

• Knowing history

• Character traits and states

• State changes

• Synapomorphies

• Homoplasy

• Character state reconstruction

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Computer Simulation http://www.life10e.com/at22.02.html

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Computer Simulation http://www.life10e.com/at22.02.html

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lab Simulation

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In the laboratory, researchers produced an unambiguous phylogeny of nine viral lineages, enhancing the mutation rate to increase variation among the lineages.

Outgroup lineageA lineage

D lineage

C lineage

E lineage

F lineage

H lineage

B lineage

G lineage

Growth in presence of mutagen

400 400 400

Generations

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Thought Simulation

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Dog Breed Geneology (imperfect record)

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http://www.macwebguru.com/2010/09/11/the-dog/

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lenski Long Term E. coli evolution 1988 ——

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lenski Long Term E. coli evolution 1988 ——

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014 16

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3: Outline

• Knowing history

• Character traits and states

• State changes

• Synapomorphies

• Homoplasy

• Character state reconstruction

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Character Traits and States

• Character: !A heritable feature of an organism. !Also known as a character trait or trait.

• Character state: !The particular form that a character takes.

• Example: !Character = heart !Character state = present/absent !Character state = # of chambers

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Character trait = fur; Character state = different colors

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Model Tree

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Character Trait: Face Character State: Happy

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 1

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014 30

Character Evolution from Tree of Life video

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Homologous characters

Characters (e.g., a face) that are inherited from a common ancestor are homologous.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3: Outline

• Knowing history

• Character traits and states

• State changes

• Synapomorphies

• Homoplasy

• Character state reconstruction

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Divergence

• Species change over time

• Known (generally) as divergence, or divergent evolution.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Character trait = fur; Character state = different colors

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Character State: Frowny

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Ancestral and Derived States

• A character with the same form as in the ancestor of a group has an ancestral state.

!

• A character with a different form as in the ancestor of a group has a derived state.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Which state is ancestral? !a) Smiley Face b) Frowny Face c) Both d) Neither

Clicker Question

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Ancestral & Derived Status is Relative

• A trait may be ancestral or derived, depending on the point of reference.

• Example: feathers are an ancestral trait for any group of modern birds.

• But in a phylogeny of all vertebrates, feathers would be a derived trait, showing the relationship between birds and their theropod dinosaur relatives.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Character Evolution Example 2

Character Trait: Face Ancestral State: Happy Derived State: Frowny

For This Clade

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3: Outline

• Knowing history

• Character traits and states

• State changes

• Synapomorphies

• Homoplasy

• Character state reconstruction

51

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Synapomorphy

• Derived traits that are shared among a group and are viewed as evidence of the common ancestry of the group are known as synapomorphies.

• The vertebral column is a synapomorphy of all vertebrates.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Synapomorphy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Synapomorphy

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Synapomorphy

• Synapomorphy = a shared derived trait

!

• Derivation: Greek !Syn ~ with (a.k.a. shared) !Apo ~ away from (a.k.a. derived) !Morphy ~ shape (a.k.a. character trait or state

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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3 45

Which traits are synapomorphies for fgh?

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8 9

A: 7,8,9 B: 4,5,6,7 C: 6

D: 6,7,8,9 E: 7

Clicker Question

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e Birds Dinos f

Synapomorphy Example

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e Birds Dinos f

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34

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Synapomorphy Example

Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e Birds Dinos f

12

3

hairmilk

mammary glandsfour-chambered heart 4

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Synapomorphy Example

Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e Birds Dinos f

12

3

hairmilk

mammary glandsfour-chambered heart 4

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Synapomorphy Example

Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e Birds Dinos f

12

34

56

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Synapomorphy Example

Mammals Reptiles

hairmilk

mammary glandsfour-chambered heart

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3: Outline

• Knowing history

• Character traits and states

• State changes

• Synapomorphies

• Homoplasy

• Character state reconstruction

62

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Homoplasy

Traits that are similar for reasons other than inheritance from a common ancestor are called homoplastic traits or homoplasies.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

Homoplasy Example

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a b c d e h g f

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Homoplasy Example

Frowny face is homoplasious

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Convergence

Similar traits can develop in unrelated groups of organisms:

Convergent evolution—independently evolved traits subjected to similar selection pressures may become superficially similar.

!

Example: the wings of bats and birds

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Figure 22.4 The Bones Are Homologous, the Wings Are Not

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Convergence

a Bats c d e Birds g f

+

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can fly

can’t fly

Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Lecture 3 Outline

• Knowing history

• Character traits and states

• State changes

• Synapomorphies

• Homoplasy

• Character state reconstruction

77

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Character State Reconstruction

Character state reconstruction methods allow one to infer history of states and state changes

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Character State Reconstruction

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Character State Reconstruction

In theory, each node could have either of the two states

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Character State Reconstruction

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Character State Reconstruction

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Character State Reconstruction

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Character State Reconstruction

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Character State Reconstruction

Which pattern is the most plausible?

+++

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Pattern 1 Pattern 2 Pattern 3

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Principle of parsimony

The Principle of Parsimony: the simplest of two (or more) competing theories is to be preferred (assuming all else is equal).

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Key Concept: Principle of parsimony

The Principle of Parsimony: the simplest of two (or more) competing theories is to be preferred (assuming all else is equal). !

Which character state pattern requires the fewest changes in states?

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Principle of parsimony

How many “steps” (state changes) required for each?

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+++Pattern 1 Pattern 2 Pattern 3

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

+

Principle of parsimony

can fly

can’t fly

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Pattern 1: How many steps?

Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

+

Principle of parsimony

can fly

can’t fly

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Pattern 1: How many steps?

Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

Mammals Reptiles

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Principle of parsimony

Note you could have more changes but seven is the minimum needed. We focus on the minimum needed.

Pattern 1: Seven Steps is the minimum needed.

can fly

can’t fly

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Principle of parsimony

Pattern 2: How many steps?

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Principle of parsimony

Pattern 2: Two steps

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Principle of parsimony

Pattern 3: How many steps?

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Principle of parsimony

Pattern 3: Six steps

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Principle of parsimony

+++

Which pattern is the most plausible?

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Pattern 1 Pattern 2 Pattern 3

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Principle of parsimony

Which pattern is the most plausible?

+++Pattern1: 7 Steps Pattern2: 2 Steps Pattern3: 6 Steps

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Principle of parsimony

+++

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Which pattern is the most plausible?

Pattern1: 7 Steps Pattern2: 2 Steps Pattern3: 6 Steps

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Principle of parsimony

Pattern 2: Flight in birds and bats arose separately and is homoplasious.

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

a Bats c d e Birds g f

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Principle of parsimony

ASSUMPTIONS …

can fly

can’t fly

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Mammals Reptiles

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Figure 22.4 The Bones Are Homologous, the Wings Are Not

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Figure 22.10 The Origin of a Sexually Selected Trait

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Figure 22.10 The Origin of a Sexually Selected Trait

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Figure 22.10 The Origin of a Sexually Selected Trait

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014 105

SIVsyk (Sykes monkeys)

HIV-1 (humans)

SIVcpz (chimpanzees)

SIVhoest (L’Hoest monkeys)SIVsun (sun-tailed monkeys)

SIVmnd (mandrills)

SIVagm (African green monkeys)SIVsm (sooty mangabeys)

HIV-2 (humans)

Common ancestor

Where did HIV Come From?

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014 106

Common ancestor

Where did HIV Come From?

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014 107

Common ancestor

Where did HIV Come From?

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014 108

Common ancestor

Where did HIV Come From?

Virus transferred from simian host to humans

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Which Came First Red or Green Fluorescent Protein

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Kinds of Phylogenetic Trees

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A B C D

A

BC

D

A B C D

Units of character divergence

Units of time

Cladogram Phylogram Chronogram

Different kinds of trees

Only the relative branching order is depicted. No meaning to branch lengths.

Branch length is proportional to the amount of character change

Branch length is proportional to time

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Next Lecture: Phylogenetic Tree Inference

What if you did not know the phylogeny and only knew the traits and the organisms? How would you infer homology, convergence, homoplasy, etc?

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Table 22.1

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

22.2 How Are Phylogenetic Trees Constructed?

The root of the tree is located between the ingroup and the outgroup.

A trait present in both ingroup and outgroup must have evolved before the ingroup and thus must be ancestral for the ingroup.

Lampreys (jawless fishes) arose before the lineage leading to other vertebrates—they are the outgroup.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

22.2 How Are Phylogenetic Trees Constructed?

Chimpanzees and mice share two derived traits—fur and mammary glands. They are synapomorphies for this group.

Keratinous scales are a synapomorphy of the crocodile, pigeon, and lizard.

Information about the synapomorphies allows construction of the tree.

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Figure 22.5 Constructing a Phylogenetic Tree

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Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014

Figure 22.5 Constructing a Phylogenetic Tree

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