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imating the reliability of a tr Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable are they?

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Page 1: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Estimating the reliability of a treeReconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable are they?

Page 2: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Reliability most of the time, ‘reliability’ refers to the topology, not to branch lengths.

reliability = probability that the members of a given clade are always members of that clade

Page 3: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Methods phylogeneticists use different methods to test the reliability of trees

• Bootstrapping• Jackknife• Permutation tests• Likelihood ratio tests (a)LRT

Page 4: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Bootstrapping bootstrapping uses random sampling with replacement to obtain properties of an estimator

Page 5: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Bootstrapping bootstrapping uses random sampling with replacement to obtain properties of an estimator

Page 6: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

x

Bootstrapping bootstrapping uses random sampling with replacement to obtain properties of an estimator

xx

xx

1000-10000 times

x

f

Page 7: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Bootstrapping in phylogenetic bootstrapping, the alignment is resampled

A A TC G CA G TT G TT C T

1ACAAG

2TCTTT

3GGGGG

4GGGGG

5GGACG

6 7TCAGA

8 9TGTTT

0ACATT

1GGGGG

4TCAGA

8TCTTT

3GGACG

6TGTTT

0ACAAG

2GGGGG

4TCTTT

9GGGGG

5

original alignment pseudo alignment

Page 8: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Bootstrapping

A B C D E F A C B D E F

original tree bootstrapped tree

+1

+1+1

+1

+1+1

+1+0

+1+1

Page 9: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

A C B D E FA C B D E F

A C B D E FA C B D E F

Bootstrapping

A B C D E F A C B D E F

original tree bootstrapped trees

0.810.75

0.87

0.95

0.800.90

0.50.47

0.450.23

Page 10: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Jackknife methods the Jackknife procedure uses random sampling without replacement to obtain properties of an estimator

Page 11: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Jackknife methods the Jackknife procedure uses random sampling without replacement to obtain properties of an estimator

Page 12: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Permutation methods Permutation tests are standard in non-parametric statistics. They reorder the data to obtain a null distribution.

Page 13: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Permutation methods Permutation tests are standard in non-parametric statistics. They reorder the data to obtain a null distribution.

N=18, x=20 N=10, x=25 Dif=5

Page 14: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Permutation methods

N=18, x=23 N=10, x=19.6 Dif=3.4

Page 15: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Permutation methods

difference

f

5% largest differences5% smallest differences

actual difference

Page 16: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Permutation methods in phylogenetics, species can be permuted within characters

A A TC G CA G TT G TT C T

1ACAAG

2TCTTT

3GGGGG

4GGGGG

5GGACG

6 7TCAGA

8 9TGTTT

0species 1

species 2

species 3

species 4

species 5

ACATT

1

A

C

AT

T

1

reshuffle

Page 17: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Permutation methods in phylogenetics, species can be permuted within characters

A A TC G CA G TT G TT C T

1ACAAG

2TCTTT

3GGGGG

4GGGGG

5GGACG

6 7TCAGA

8 9TGTTT

0species 1

species 2

species 3

species 4

species 5

A

C

AT

T

1

AC

A

AG2

ACAAG

2

reshuffle

Page 18: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Permutation methods in phylogenetics, species can be permuted within characters

A A TC G CA G TT G TT C T

1ACAAG

2TCTTT

3GGGGG

4GGGGG

5GGACG

6 7TCAGA

8 9TGTTT

0species 1

species 2

species 3

species 4

species 5

A

C

AT

T

1

AC

A

AG2

A

TG

CG

TG T

CT

T

C

TT

T

3GGGGG

4GGGGG

5

G

GA

C

G

6 7TCA

GA

8 9

T

G

TT

T0

Page 19: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Likelihood ratio tests

A B C D E F G H I J K L

X (ABCDEF) Y (GHI)

W (J)Z (KL)

Page 20: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Likelihood ratio tests standard likelihood tests compare trees with and without the branch

X (ABCDEF) Y (GHI)

W (J)Z (KL)

X Y

WZ

Likelihood = L1 Likelihood = L0

probability that branch exists = 2 * [ln L1 – ln L0]

Page 21: Estimating the reliability of a tree Reconstructed phylogenetic trees are almost certainly wrong. They are estimates of the true tree. But how reliable

Approximate likelihood ratio test aLRT is fast, accurate and powerful

X Y

WZ

Likelihood = L1

Likelihood = L2

approximate probability that branch exists = 2 * [ln L1 – ln L2]

X Z

WY

Likelihood = L3

X Z

YW