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Effects of ancestral character
    estimation on methods for
 estimating rates of evolution in
discrete morphological characters

  Graeme T. Lloyd, Steve C. Wang and
        Stephen L. Brusatte
G. G. Simpson




          1902 - 1984
T. S. Westoll




            1912 - 1995
Westoll and lungfish




                   Westoll 1949
Simpson modification
Textbook example
Other discrete character studies
Derstler 1982                         Forey 1988




                Ruta et al. 2006   Brusatte et al. 2008
Translating Westoll’s method
Evolution is phylogenetic


    Phyletic       Phylogenetic
Evolution is phylogenetic




       Use cladogram
Zero duration branch problem
Zero duration branch problem




Use Ruta et al. dating technique
Uncertainty in dating

            Time
Uncertainty in dating




Randomly choose dates and repeat
Uncertainty in optimisation
Uncertainty in optimisation




Apply ACCTRAN and DELTRAN
Uncertainty in topology
Uncertainty in topology




   Apply to multiple MPTs
Specimen completeness bias

Many elements preserved   Few elements preserved




Many changes observed      Few changes observed
Specimen completeness bias




Use Wagner’s patristic dissimilarity
Lacks null/alt. hypotheses




    H0             Halt
Lacks null/alt. hypotheses




Apply equal rates as null hypothesis
Data
Lungfish
Lungfish supermatrix
                                 91 characters




                       86 taxa
Results
Rate over time test (ACCTRAN)
Rate over time test (DELTRAN)
Likelihood estimation of ancestral states
Parsimony problems

  DELTRAN           ACCTRAN




                ?

                ?
                              Change
                               early
                ?


       Change
        late
Likelihood




                     ?

         ?           ?
             ?
                     ?
                 ?
Likelihood




                           ?

             ?             ?
                   ?
                           ?
      No changes
                       ?
Rate over time test (Likelihood)
Rate over time test (Likelihood)
Terminal vs internal branches
Branch length comparison
Branch rate comparison
Branch changes comparison
Branch duration comparison
Branch length comparison
Internal length comparison


  Devonian


             post-Devonian
Terminal length comparison




 Devonian

                 post-Devonian
Conclusions
Acknowledgements, code, data




          Roger Benson



 http://www.graemetlloyd.com/

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Effects of ancestral character estimation on methods for estimating rates of evolution in discrete morphological characters