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Flexible Modeling of
Morphological Characters for
Phylogenetic Inference
April Wright
Will Pett
Tracy Heath
@wrightingapril
Challenges of Modeling
Morphological Data
Common meaning is not carried across characters
Challenges of Modeling
Morphological Data
Nucleotides do carry common meaning across
characters
ACTGAGTCGAATGCTGGGAGCTA
Challenges of Modeling
Morphological Data
Nucleotides do carry common meaning across
characters
A G
C T
Challenges of Modeling
Morphological Data
Common meaning is not carried across characters
01110112011131102210
Challenges of Modeling
Morphological Data
Common meaning is not carried across characters
01110112011131102210
Challenges of Modeling
Morphological Data
Common meaning is not carried across characters
01110112011131102210
0
0 1
Challenges of Modeling
Morphological Data
Common meaning is not carried across characters
01110112011131102210
0
0 1
0
0 1
1
Challenges of Modeling
Morphological Data
Image by Chris Smith
Challenges of Modeling
Morphological Data
Image by Chris Smith
Challenges of Modeling
Morphological Data
Image by Chris Smith
Challenges of Modeling
Morphological Data
0 1
Common meaning is not carried across characters
The Mk Model
(Lewis, 2001)
Challenges of Modeling
Morphological Data
0 1
2 3
Common meaning is not carried across characters
The Mk Model
Symmetric Beta Prior
↵ =
Figure from Wright et al. 2016,
model implemented in MrBayes,
Ronquist and Huelsenbeck 2003
Percentage of transitions
0 to 1
1 to 0
Symmetric Beta Prior
↵ =
Figure from Wright et al. 2016,
model implemented in MrBayes,
Ronquist and Huelsenbeck 2003
Percentage of transitions
0 to 1
1 to 0
Ant Morphological Data
Image by Steve Shattuck
Ants have a rich fossil record, including many stem lineages,
as well as many scored specimens from extant lineages
Ant Morphological Data
Image by Steve Shattuck
Presence of wasplike
features that
are lost after
divergence of
Sphecomyrminae
from the rest of
the ants
1 0
Ant Morphological Data
Image by Steve Shattuck
Presence of wasplike
features that
are lost after
divergence of
Sphecomyrminae
from the rest of
the ants
1 0
Presence of ant
apomorphies
0 1
Ant Morphological Data
Image by Steve Shattuck
Presence of wasplike
features that
are lost after
divergence of
Sphecomyrminae
from the rest of
the ants
1 0
Presence of ant
apomorphies
0 1
Strong violation of equal
transition rates model
Ant Morphological Data
Image by Steve Shattuck
• Recoded characters from Barden
and Grimaldi 2016; Keller 2011
• 164 characters, 135 taxa
Ant Morphological Data
Image by Steve Shattuck
• Recoded characters from Barden
and Grimaldi 2016; Keller 2011
• 164 characters, 135 taxa
byfir001
Symmetric Beta Prior
↵ =
Figure from Wright et al. 2016,
model implemented in MrBayes,
Ronquist and Huelsenbeck 2003
Percentage of transitions
0 to 1
1 to 0
Asymmetric Beta Prior
↵ 6=
Asymmetric Beta Prior
↵ 6=
Asymmetric Beta Prior
↵ 6=
0 1
1 0
Asymmetric Beta Prior
↵ 6=
0 1
1 0
0 1
1 0
Asymmetric Beta Prior
↵ 6=
0 1
1 0
0 1
1 0
Asymmetric Beta Prior
↵ 6=
0 1
1 0
0 1
1 0
0 1
1 0
Asymmetric Beta Prior
↵ 6=
0 1
1 0
0 1
1 0
0 1
1 0
0 1
1 0
Symmetric Beta Prior
↵ =
This is a special case of the asymmetrical Beta
Percentage of transitions
0 to 1
1 to 0
Asymmetric Beta Prior
We also implemented a multistate generalization
in which state frequencies are drawn from a Dirichlet
prior of k dimensions
Ant Morphological Data
• Recoded from Barden and
Grimaldi 2016; Keller 2011
• 164 characters, 135 taxa
Image by Steve Shattuck
byfir001
Ant Morphological Data
• Estimated phylogeny
from binary data sets
using discrete Beta
model
Image by Steve Shattuck
byfir001
Ant Morphological Data
• Estimated phylogeny
from binary data sets
using discrete Beta
model
• Phylogeny using
binary + multistate
data with generalized
Beta model
Image by Steve Shattuck
byfir001
Asymmetric Beta Prior
↵ 6=
0 1
1 0
0 1
1 0
Results: Asymmetric Beta
Model
• Does using the Asymmetric Beta Model improve
model fit for this dataset?
Asymmetric Beta Model
Improves Model Fit
Asymmetric Beta Model
Improves Model Fit
Mk model
Amount of Heterogeneity Is
Detectable With Model Fitting
State Frequencies Biased
Toward 0 Character
State Frequencies Biased
Toward 0 Character
More 0 1 transitions in dataset
Results: Asymmetric
Multistate Model
• Does using the asymmetric multistate model
improve model fit for this dataset?
Asymmetric Multistate Model
Improves Model Fit
Asymmetric Multistate Model
Improves Model Fit
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
Improved Models Improve
Phylogeny
PP > .9
PP > .5
PP > .1
Conclusions
• Allowing for transition rate asymmetry in both
binary and multistate characters is very helpful in
this dataset
Conclusions
• Allowing for transition rate asymmetry in both
binary and multistate characters is very helpful in
this dataset
• The number of transition symmetry categories is
detectable with Bayes factor model fitting
Conclusions
• Allowing for transition rate asymmetry in both
binary and multistate characters is very helpful in
this dataset
• The number of transition symmetry categories is
detectable with Bayes factor model fitting
• Resolving ant phylogenetic relationships remains
challenging, but hopeful
Thank You
The Heath Lab
Tracy Heath
Walker Pett
Jordan Satler
Kevin Quinteros
Wade Dismukes
Roberta Graboski Mendes
Andre Flores
Patrick Mendoza
Helpful
Discussions
Corrie Moreau
Paul Lewis
Benjamin Blanchard
Philip Barden
Mark Holder
DEB-1556615, DEB-1256993
DBI-1612858
revbayes.github.io
Ant Phylogeny Remains
Challenging
“There are only so many ways to make an ant”
-Corrie Moreau
Wright Lab
Ant Phylogeny Is
Challenging
“There are only so many ways to make an ant”
-Corrie Moreau
What means for ants?
Results: Asymmetric Beta
Model
Results: Asymmetric Beta
Model

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