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Fossils-only tip-dating of 
deinonychosaurian 
theropods 
April M. Wright 
Nicholas J. Matzke 
Graeme T. Lloyd 
David W. Bapst
www.slideshare.net/wrightaprilm/wright-svp2014
Can we use fossils-as-tips methods to co-estimate 
phylogeny and divergence dates in 
the absence of molecular data?
Theropods as a test case for tip-dating 
● Controversial 
● Implications for the evolution of flight 
● Well-documented ages and relationships 
among taxa in the tree
Placement of Archaeopteryx
Placement of Archaeopteryx
Placement of Archaeopteryx
A paleontological question
A paleontological question
A paleontological question
A paleontological question 
Archaeopteryx as 
a calibration 
node
Applying tip-dating methods 
● Used Xu et al. matrix 
● 89 taxa, 389 morphological characters 
○ Missing data ranges between ~5% and ~85% per 
taxon 
○ 363 characters from Hu et al. 2009, 10 added by Xu 
et al. 
○ Mix of cranial and post-cranial characters
Applying tip-dating methods 
● Dates associated with every terminal in the 
matrix 
○ Both a minimum and a maximum age
Applying tip-dating methods 
● Dates associated with every terminal in the 
matrix 
○ Both a minimum and a maximum age 
● BEAST2 and MrBayes tip dating 
○ Both methods use the dates of the terminal taxa 
instead of calibration points 
○ Both use Lewis’ 2001 Mk model
Applying tip-dating methods 
● BEAST2 
○ Birth-death serial sampling prior: Allows for 
speciation and extinction in tree for data sampled 
from multiple time-points 
● MrBayes 
○ Uniform prior on clock trees 
○ Ingroup date calibration
Applying tip-dating methods 
● BEAST2 
○ More explicit process-driven model 
● MrBayes 
○ More vague with respect to biology and taphonomy
} Aves 
}Deinonychosauria 
200 175 150 125 100 75 50 25
} Aves 
}Deinonychosauria 
200 175 150 125 100 75 50 25
} Aves 
}Deinonychosauria 
200 175 150 125 100 75 50 25
Placement of Archaeopteryx
MrBayes 
● We were unable to obtain a topology with 
dates in which we had confidence 
○ Well-known clades being broken up 
○ Unrealistic dates that contradict the weight of 
existing evidence
MrBayes 
Xu et al. 2006
MrBayes
Conclusions 
● Tip-dating is a viable method for estimating 
divergence dates 
○ Tip-dating can be performed with fossil-only 
datasets 
● BEAST returns a topology and dates that 
correspond well to known information about 
deinonychosaurian and avian relationships
Thanks! 
● Participants 
● Hillis Lab 
Group 
● Nadia 
Fröbisch, 
Jonathan 
Bloch, Anjali 
Goswami

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April Wright - SVP 2014