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New methodologies for the use of 
cladistic-type matrices to measure 
morphological disparity and 
evolutionary rate 
@GraemeTLloyd
Acknowledgements 
Matt 
Friedman 
Liam 
Revell 
Mark 
Bell 
Peter 
Smits 
Steve 
Brusatte 
Roger 
Benson 
Steve 
Wang 
Rich 
Fitzjohn
Cladistic-type data 
- Discrete morphological data
Cladistic-type data 
- Discrete morphological data 
- Limited to 32 states (often 
less)
Cladistic-type data 
- Discrete morphological data 
- Limited to 32 states (often 
less) 
- Frequently non-Euclidean
Cladistic-type data 
- Discrete morphological data 
- Limited to 32 states (often 
less) 
- Frequently non-Euclidean 
- Missing data common
Acladistic analyses 
Disparity Rates
Acladistic analyses 
Disparity Rates 
Common Rare
Acladistic analyses 
Disparity Rates 
Common 
No models 
Rare 
Simple models
Acladistic analyses 
Disparity Rates 
Common 
No models 
Single approach (GED) 
Rare 
Simple models 
N approaches ≈ N studies
Acladistic analyses 
Disparity Rates 
Common 
No models 
Single approach (GED) 
Rare 
Simple models 
N approaches ≈ N studies 
Time series an issue
Claddis 
github.com/graemetlloyd/Claddis
Disparity
Toljagic 
and 
Butler 
2013 
Disparity studies 
Brusatte et al 
2008 
Thorne 
et al 
2011 
Butler et al. 2011
Cladistic disparity 
Cladistic 
matrix 
Distance 
matrix 
Ordination ‘Morphospace’
Cladistic disparity 
Cladistic 
matrix 
Distance 
matrix 
Ordination ‘Morphospace’ 
Distance 
metric
Desiderata 
An ideal distance metric should:
Desiderata 
An ideal distance metric should: 
1. have high fidelity
Desiderata 
An ideal distance metric should: 
1. have high fidelity 
2. be normally distributed
Desiderata 
An ideal distance metric should: 
1. have high fidelity 
2. be normally distributed 
3. be Euclidean
Desiderata 
An ideal distance metric should: 
1. have high fidelity 
2. be normally distributed 
3. be Euclidean 
4. be calculable
Desiderata 
An ideal distance metric should: 
1. have high fidelity 
2. be normally distributed 
3. be Euclidean 
4. be calculable 
5. be easily visualised
Generalised Euclidean Distance 
Wills 2001
Generalised Euclidean Distance 
But: Sijk is incalculable if k values for i or j (or both) are missing 
Wills 2001
Generalised Euclidean Distance 
But: Sijk is incalculable if k values for i or j (or both) are missing 
Wills 2001
Alternate distances 
GED
Alternate distances 
Raw GED
Alternate distances 
Raw GED 
Gower
Alternate distances 
Raw GED 
Gower 
MOD
Simulations 
Input 
20 taxa 
50 binary characters 
0-80% missing data
Simulations 
Input Output 
20 taxa 
50 binary characters 
0-80% missing data 
N taxa retained 
Variance of first two PCA axes 
Shapiro-Wilk test 
Mantel test
Calculable 
Raw GED 
Gower 
MOD 
Incompleteness 
100% 
% taxa retained 
0% 80% 
0%
Visualisation 
Raw GED 
Gower 
MOD 
Incompleteness 
45% 
% variance axes 1 & 2 
0% 80% 
15%
Normalcy 
Raw GED 
Gower 
MOD 
Incompleteness 
1.00 
Shapiro-Wilk test 
0% 80% 
0.75
Fidelity 
Raw GED 
Gower 
MOD 
Incompleteness 
+30% 
Correlation 
0% 80% 
-30%
Fidelity 
Gower 
Raw 
GED 
MOD 
Incompleteness 
100% 
% highest fidelity 
0% 80% 
0%
% missing data 
Incompleteness 
30 
N data sets 
0% 80% 
0
Rates
Rate studies 
Derstler 1982 Forey 1988 
Ruta et al 2006 Brusatte et al 2008
Rate calculation 
Rate = N changes / 
Δt × Completeness
Null hypothesis 
H0 = equal rates
Alternate hypothesis 
Halt = +
Lungfish 
Westoll 1949
Lungfish 
Devonian 
high rates 
Lloyd et al 2012
Lungfish 
post-Devonian 
low rates 
Lloyd et al 2012
Parsimony problem 
DELTRAN ACCTRAN 
? 
? 
? 
Change 
early 
Change 
late
Parsimony problem 
Lloyd et al 2012
Parsimony problem 
Lloyd et al 2012
Parsimony problem 
? 
? 
? 
? 
? 
?
Parsimony problem 
? 
? 
? 
? 
? 
? 
? 
? 
? 
? 
? 
? No changes
Internal vs. terminal 
Rate 
>
Internal vs. terminal 
Changes 
≈
Internal vs. terminal 
Duration 
<
Internal vs. terminal 
Solution
Rates revisited 
high rates 
low rates 
Brusatte et al 2014
Rates revisited 
high rates 
low rates 
Brusatte et al 2014
Time series problems
Toljagic 
and 
Butler 
2013 
Disparity time series 
Brusatte et al 
2008 
Thorne 
et al 
2011 
Butler et al. 2011
Toljagic 
and 
Butler 
2013 
Disparity time series 
4 time bins 4 time bins 
Brusatte et al 
2008 
Thorne 
et al 
2011 
14 time bins 2 time bins 
Butler et al. 2011
Rate time series 
Lloyd et al 2012 
Ruta et al 
2006 
Branch-binning No completeness
Rate time series 
N changes | Δt | Completeness
Conclusions 
Raw GED 
Gower 
MOD
Conclusions 
Raw GED 
Gower 
MOD 
PCO 1 
? 
PCO 2
Conclusions 
Raw GED 
Gower 
MOD 
PCO 1 
? 
PCO 2
Conclusions 
Raw GED 
Gower 
MOD 
Rate 
? 
? 
t 
PCO 1 
PCO 2

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New methodologies for the use of cladistic-type matrices to measure morphological disparity and evolutionary rate

Notes de l'éditeur

  1. Fidelity (distances in ordination = true distances) Normality (no outliers) Distances should be Euclidean (low/no negative eigenvalues) High variance on first two ordination axes (visualisation) Retain as many taxa as possible (calculable distances)
  2. Fidelity (distances in ordination = true distances) Normality (no outliers) Distances should be Euclidean (low/no negative eigenvalues) High variance on first two ordination axes (visualisation) Retain as many taxa as possible (calculable distances)
  3. Fidelity (distances in ordination = true distances) Normality (no outliers) Distances should be Euclidean (low/no negative eigenvalues) High variance on first two ordination axes (visualisation) Retain as many taxa as possible (calculable distances)
  4. Fidelity (distances in ordination = true distances) Normality (no outliers) Distances should be Euclidean (low/no negative eigenvalues) High variance on first two ordination axes (visualisation) Retain as many taxa as possible (calculable distances)
  5. Fidelity (distances in ordination = true distances) Normality (no outliers) Distances should be Euclidean (low/no negative eigenvalues) High variance on first two ordination axes (visualisation) Retain as many taxa as possible (calculable distances)
  6. Fidelity (distances in ordination = true distances) Normality (no outliers) Distances should be Euclidean (low/no negative eigenvalues) High variance on first two ordination axes (visualisation) Retain as many taxa as possible (calculable distances)