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A general model for continuous character
                            evolution

                            Carl Boettiger

                               UC Davis


                            June 20, 2011




Carl Boettiger, UC Davis          Release of Constraint   1/37
A Key Innovation in Jaw Morphology?




Carl Boettiger, UC Davis   Release of Constraint   2/37
A long time ago. . .




Carl Boettiger, UC Davis            Release of Constraint   3/37
This fish




Carl Boettiger, UC Davis      Release of Constraint   4/37
Got a good idea




      Westneat et. al. (2005)




Carl Boettiger, UC Davis               Release of Constraint   5/37
and did this:




Carl Boettiger, UC Davis   Release of Constraint   6/37
and did this:




Carl Boettiger, UC Davis   Release of Constraint   6/37
Parrotfish with the intramandibular joint:




Carl Boettiger, UC Davis   Release of Constraint   7/37
Parrotfish with the intramandibular joint:
              have over 6 times greater disparity
              in their jaw opening lever ratio
              have over 3 times greater disparity
              in their closing lever ratio
              comparable variation in protrusion
              and they are a younger clade
              relative to other parrotfish

      (corrected for body mass)

                                                                 Price et. al. (2010)




Carl Boettiger, UC Davis                 Release of Constraint                          7/37
A Key Innovation* in Jaw Morphology?




                       * With respect to the resulting diversity of morphology,
                                                  rather than species diversity



Carl Boettiger, UC Davis                     Release of Constraint                8/37
One group remains under
              selective constraint
              One with innovation can
              diversify morphology




Carl Boettiger, UC Davis        Release of Constraint   9/37
This is a nice question for comparative
                phylogenetic models. . .




Carl Boettiger, UC Davis   Release of Constraint   10/37
We don’t have that model.




Carl Boettiger, UC Davis               Release of Constraint   11/37
I have that model




Carl Boettiger, UC Davis           Release of Constraint   12/37
Models for Continuous Character
                           Evolution




Carl Boettiger, UC Davis       Release of Constraint   13/37
A Phylogeny of Phylogenetic Models

     BM                Brownie                            ouch     OU
                           2006



                                                            2004


                           1999




                                            1997




                                                     1985




Carl Boettiger, UC Davis          Release of Constraint                 14/37
Something is missing

                    Brownie:
   Differing rates of diversification
                                                                  ouch:
                                                    Differing selective optima




                           Where’s the link? Can it be both?




Carl Boettiger, UC Davis                  Release of Constraint                  15/37
Some comparative physiology: Brownie


                              dX          = σ × dBt
                           trait change    rate        Brownian




Carl Boettiger, UC Davis                   Release of Constraint   16/37
Some comparative physiology: Brownie


                              dX          = σ × dBt
                           trait change    rate        Brownian




Carl Boettiger, UC Davis                   Release of Constraint   16/37
Some comparative physiology: ouch


                           dX = α(   θ     −X)dt + σdBt
                                 optimum




Carl Boettiger, UC Davis                 Release of Constraint   17/37
Some comparative physiology: ouch


                           dX = α(   θ     −X)dt + σdBt
                                 optimum




Carl Boettiger, UC Davis                 Release of Constraint   17/37
How about:



                           dX =      α (θ − X)dt + σdBt
                                  selection




Carl Boettiger, UC Davis                      Release of Constraint   18/37
How about:



                           dX =      α (θ − X)dt + σdBt
                                  selection




Carl Boettiger, UC Davis                      Release of Constraint   18/37
Combine both:


                    dX = α(   θ      −X)dt +                  σ     dBt
                           optimum                   divers. rate




Carl Boettiger, UC Davis              Release of Constraint               19/37
Combine both:


                    dX = α(   θ      −X)dt +                  σ     dBt
                           optimum                   divers. rate




Carl Boettiger, UC Davis              Release of Constraint               19/37
Many other combinations:


                       dX =     α (       θ       −X)dt + σdBt
                              selection optimum




Carl Boettiger, UC Davis                   Release of Constraint   20/37
Many other combinations:


                       dX =     α (       θ       −X)dt + σdBt
                              selection optimum




Carl Boettiger, UC Davis                   Release of Constraint   20/37
One model to rule them all?


               dX =          α (       θ       −X)dt +                     σ          dBt
                           selection optimum                           divers. rate




Carl Boettiger, UC Davis                       Release of Constraint                        21/37
One model to rule them all?


               dX =          α (       θ       −X)dt +                     σ          dBt
                           selection optimum                           divers. rate




Carl Boettiger, UC Davis                       Release of Constraint                        21/37
But do you have the power to wield it?




Carl Boettiger, UC Davis   Release of Constraint   22/37
But do you have the power to wield it?



     Do you have . . . the complete
      time-calibrated tree of life?




Carl Boettiger, UC Davis              Release of Constraint   22/37
But do you have the power to wield it?



     Do you have . . . the complete             Or is your data a hobbit?
      time-calibrated tree of life?




Carl Boettiger, UC Davis              Release of Constraint                 22/37
A dangerous path




Carl Boettiger, UC Davis   Release of Constraint   23/37
A dangerous path
     Ridges so flat and long you may
     be stuck forever




Carl Boettiger, UC Davis   Release of Constraint   23/37
A dangerous path
     Ridges so flat and long you may
     be stuck forever


              Rugged likelihood surfaces such that
              common ML algorithms (Nelder-Mead,
              L-BFGS-B), will fail




Carl Boettiger, UC Davis        Release of Constraint   23/37
A dangerous path
     Ridges so flat and long you may
     be stuck forever


              Rugged likelihood surfaces such that
              common ML algorithms (Nelder-Mead,
              L-BFGS-B), will fail


              AIC, familiar guide to model choice,
              is slimy and treacherous



Carl Boettiger, UC Davis          Release of Constraint   23/37
A dangerous path
     Ridges so flat and long you may
     be stuck forever


              Rugged likelihood surfaces such that
              common ML algorithms (Nelder-Mead,
              L-BFGS-B), will fail


              AIC, familiar guide to model choice,
              is slimy and treacherous



Carl Boettiger, UC Davis          Release of Constraint   23/37
Likelihood ridges




              α ∝ σ 2 ridge-line




Carl Boettiger, UC Davis           Release of Constraint   24/37
Likelihood ridges




              α ∝ σ 2 ridge-line
              small or deeply branching
              phylogenies




Carl Boettiger, UC Davis      Release of Constraint   24/37
Likelihood ridges




              α ∝ σ 2 ridge-line
              small or deeply branching
              phylogenies
              Closely nested regime paintings




Carl Boettiger, UC Davis      Release of Constraint   24/37
Survival Strategies
on rugged likelihood surfaces



              Sub-models




Carl Boettiger, UC Davis   Release of Constraint   25/37
Survival Strategies
on rugged likelihood surfaces



              Sub-models
              Simulated annealing (if max-likelihood)




Carl Boettiger, UC Davis          Release of Constraint   25/37
Survival Strategies
on rugged likelihood surfaces



              Sub-models
              Simulated annealing (if max-likelihood)
              MCMC → posterior distributions (in
              Bayesian mode)




Carl Boettiger, UC Davis          Release of Constraint   25/37
Survival Strategies
on rugged likelihood surfaces



              Sub-models
              Simulated annealing (if max-likelihood)
              MCMC → posterior distributions (in
              Bayesian mode)
              PMC for power estimates & model choice*




Carl Boettiger, UC Davis         Release of Constraint   25/37
Survival Strategies
on rugged likelihood surfaces



              Sub-models
              Simulated annealing (if max-likelihood)
              MCMC → posterior distributions (in
              Bayesian mode)
              PMC for power estimates & model choice*
              (“A simulation analysis in a box” for your study)

                                                         * Boettiger et al. 2011 in review


Carl Boettiger, UC Davis                       Release of Constraint                         25/37
Returning to our opening example. . .




Carl Boettiger, UC Davis   Release of Constraint   26/37
Returning to our opening example. . .




Carl Boettiger, UC Davis   Release of Constraint   26/37
Can a key innovation release selective constraint?


  Differing trait diversification       Differing strengths of stabilizing
  rates, or                            selection?




Carl Boettiger, UC Davis           Release of Constraint               27/37
Differing Trait Diversification Rate Model




Carl Boettiger, UC Davis   Release of Constraint   28/37
Differing Stabilizing Selection Strength




Carl Boettiger, UC Davis   Release of Constraint   29/37
So which is the better model?



          Differing Diversification
          Log Likelihood: -94
          Parameters: 4
          Differing Selection Log
          Likelihood: -92
          Parameters: 4




Carl Boettiger, UC Davis             Release of Constraint   30/37
So which is the better model?



          Differing Diversification
          Log Likelihood: -94
          Parameters: 4
          Differing Selection Log
          Likelihood: -92
          Parameters: 4




Carl Boettiger, UC Davis             Release of Constraint   30/37
So which is the better model?



          Differing Diversification
          Log Likelihood: -94
          Parameters: 4
          Differing Selection Log
          Likelihood: -92
          Parameters: 4




      Not enough power for this comparison in this trait.

Carl Boettiger, UC Davis             Release of Constraint   30/37
So which is the better model?



          Differing Diversification
          Log Likelihood: -94
          Parameters: 4
          Differing Selection Log
          Likelihood: -92
          Parameters: 4




      Not enough power for this comparison in this trait.
      . . . must try harder. . .
Carl Boettiger, UC Davis             Release of Constraint   30/37
A slightly simpler model comparison


  Differing trait diversification       Differing strengths of stabilizing
  rates, or                            selection?




Carl Boettiger, UC Davis           Release of Constraint               31/37
Differing Stabilizing Selection Strength




                             Differing Trait Diversification Rate




Carl Boettiger, UC Davis                      Release of Constraint   32/37
Comparison




Carl Boettiger, UC Davis   Release of Constraint   33/37
If we prefer to be Bayesian: MCMC




              These are posteriors, not likelihood bootstraps. Requires
              priors, etc.
              But, looks pretty much the same.


Carl Boettiger, UC Davis                Release of Constraint             34/37
Conclusions

         1    A new phylogenetic method




Carl Boettiger, UC Davis             Release of Constraint   35/37
Conclusions

         1    A new phylogenetic method



         2    Complex models are dangerous with inadequate power




Carl Boettiger, UC Davis             Release of Constraint         35/37
Conclusions

         1    A new phylogenetic method



         2    Complex models are dangerous with inadequate power


         3    Have methods to quantify power and uncertainty.




Carl Boettiger, UC Davis               Release of Constraint       35/37
Conclusions

         1    A new phylogenetic method



         2    Complex models are dangerous with inadequate power


         3    Have methods to quantify power and uncertainty.



         4    Hobbit-sized swords: Smaller sub-models for smaller data




Carl Boettiger, UC Davis               Release of Constraint             35/37
Acknowledgements

      Peter Wainwright
      Wainwright Lab
      Graham Coop
      Peter Ralph
      Funding:

                                      Download the development version now:
                                      https://github.com/cboettig/wrightscape




                           Slides and more our on new Lab Blog:
                            http://wainwrightlab.wordpress.com
Carl Boettiger, UC Davis                    Release of Constraint          36/37
An Addendum on Reproducible Research
              Slides on http://wainwrightlab.wordpress.com
              See meta-data, script, version of code, data, recent
              changes. . .




Carl Boettiger, UC Davis                Release of Constraint        37/37

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A general model of continuous character evolution

  • 1. A general model for continuous character evolution Carl Boettiger UC Davis June 20, 2011 Carl Boettiger, UC Davis Release of Constraint 1/37
  • 2. A Key Innovation in Jaw Morphology? Carl Boettiger, UC Davis Release of Constraint 2/37
  • 3. A long time ago. . . Carl Boettiger, UC Davis Release of Constraint 3/37
  • 4. This fish Carl Boettiger, UC Davis Release of Constraint 4/37
  • 5. Got a good idea Westneat et. al. (2005) Carl Boettiger, UC Davis Release of Constraint 5/37
  • 6. and did this: Carl Boettiger, UC Davis Release of Constraint 6/37
  • 7. and did this: Carl Boettiger, UC Davis Release of Constraint 6/37
  • 8. Parrotfish with the intramandibular joint: Carl Boettiger, UC Davis Release of Constraint 7/37
  • 9. Parrotfish with the intramandibular joint: have over 6 times greater disparity in their jaw opening lever ratio have over 3 times greater disparity in their closing lever ratio comparable variation in protrusion and they are a younger clade relative to other parrotfish (corrected for body mass) Price et. al. (2010) Carl Boettiger, UC Davis Release of Constraint 7/37
  • 10. A Key Innovation* in Jaw Morphology? * With respect to the resulting diversity of morphology, rather than species diversity Carl Boettiger, UC Davis Release of Constraint 8/37
  • 11. One group remains under selective constraint One with innovation can diversify morphology Carl Boettiger, UC Davis Release of Constraint 9/37
  • 12. This is a nice question for comparative phylogenetic models. . . Carl Boettiger, UC Davis Release of Constraint 10/37
  • 13. We don’t have that model. Carl Boettiger, UC Davis Release of Constraint 11/37
  • 14. I have that model Carl Boettiger, UC Davis Release of Constraint 12/37
  • 15. Models for Continuous Character Evolution Carl Boettiger, UC Davis Release of Constraint 13/37
  • 16. A Phylogeny of Phylogenetic Models BM Brownie ouch OU 2006 2004 1999 1997 1985 Carl Boettiger, UC Davis Release of Constraint 14/37
  • 17. Something is missing Brownie: Differing rates of diversification ouch: Differing selective optima Where’s the link? Can it be both? Carl Boettiger, UC Davis Release of Constraint 15/37
  • 18. Some comparative physiology: Brownie dX = σ × dBt trait change rate Brownian Carl Boettiger, UC Davis Release of Constraint 16/37
  • 19. Some comparative physiology: Brownie dX = σ × dBt trait change rate Brownian Carl Boettiger, UC Davis Release of Constraint 16/37
  • 20. Some comparative physiology: ouch dX = α( θ −X)dt + σdBt optimum Carl Boettiger, UC Davis Release of Constraint 17/37
  • 21. Some comparative physiology: ouch dX = α( θ −X)dt + σdBt optimum Carl Boettiger, UC Davis Release of Constraint 17/37
  • 22. How about: dX = α (θ − X)dt + σdBt selection Carl Boettiger, UC Davis Release of Constraint 18/37
  • 23. How about: dX = α (θ − X)dt + σdBt selection Carl Boettiger, UC Davis Release of Constraint 18/37
  • 24. Combine both: dX = α( θ −X)dt + σ dBt optimum divers. rate Carl Boettiger, UC Davis Release of Constraint 19/37
  • 25. Combine both: dX = α( θ −X)dt + σ dBt optimum divers. rate Carl Boettiger, UC Davis Release of Constraint 19/37
  • 26. Many other combinations: dX = α ( θ −X)dt + σdBt selection optimum Carl Boettiger, UC Davis Release of Constraint 20/37
  • 27. Many other combinations: dX = α ( θ −X)dt + σdBt selection optimum Carl Boettiger, UC Davis Release of Constraint 20/37
  • 28. One model to rule them all? dX = α ( θ −X)dt + σ dBt selection optimum divers. rate Carl Boettiger, UC Davis Release of Constraint 21/37
  • 29. One model to rule them all? dX = α ( θ −X)dt + σ dBt selection optimum divers. rate Carl Boettiger, UC Davis Release of Constraint 21/37
  • 30. But do you have the power to wield it? Carl Boettiger, UC Davis Release of Constraint 22/37
  • 31. But do you have the power to wield it? Do you have . . . the complete time-calibrated tree of life? Carl Boettiger, UC Davis Release of Constraint 22/37
  • 32. But do you have the power to wield it? Do you have . . . the complete Or is your data a hobbit? time-calibrated tree of life? Carl Boettiger, UC Davis Release of Constraint 22/37
  • 33. A dangerous path Carl Boettiger, UC Davis Release of Constraint 23/37
  • 34. A dangerous path Ridges so flat and long you may be stuck forever Carl Boettiger, UC Davis Release of Constraint 23/37
  • 35. A dangerous path Ridges so flat and long you may be stuck forever Rugged likelihood surfaces such that common ML algorithms (Nelder-Mead, L-BFGS-B), will fail Carl Boettiger, UC Davis Release of Constraint 23/37
  • 36. A dangerous path Ridges so flat and long you may be stuck forever Rugged likelihood surfaces such that common ML algorithms (Nelder-Mead, L-BFGS-B), will fail AIC, familiar guide to model choice, is slimy and treacherous Carl Boettiger, UC Davis Release of Constraint 23/37
  • 37. A dangerous path Ridges so flat and long you may be stuck forever Rugged likelihood surfaces such that common ML algorithms (Nelder-Mead, L-BFGS-B), will fail AIC, familiar guide to model choice, is slimy and treacherous Carl Boettiger, UC Davis Release of Constraint 23/37
  • 38. Likelihood ridges α ∝ σ 2 ridge-line Carl Boettiger, UC Davis Release of Constraint 24/37
  • 39. Likelihood ridges α ∝ σ 2 ridge-line small or deeply branching phylogenies Carl Boettiger, UC Davis Release of Constraint 24/37
  • 40. Likelihood ridges α ∝ σ 2 ridge-line small or deeply branching phylogenies Closely nested regime paintings Carl Boettiger, UC Davis Release of Constraint 24/37
  • 41. Survival Strategies on rugged likelihood surfaces Sub-models Carl Boettiger, UC Davis Release of Constraint 25/37
  • 42. Survival Strategies on rugged likelihood surfaces Sub-models Simulated annealing (if max-likelihood) Carl Boettiger, UC Davis Release of Constraint 25/37
  • 43. Survival Strategies on rugged likelihood surfaces Sub-models Simulated annealing (if max-likelihood) MCMC → posterior distributions (in Bayesian mode) Carl Boettiger, UC Davis Release of Constraint 25/37
  • 44. Survival Strategies on rugged likelihood surfaces Sub-models Simulated annealing (if max-likelihood) MCMC → posterior distributions (in Bayesian mode) PMC for power estimates & model choice* Carl Boettiger, UC Davis Release of Constraint 25/37
  • 45. Survival Strategies on rugged likelihood surfaces Sub-models Simulated annealing (if max-likelihood) MCMC → posterior distributions (in Bayesian mode) PMC for power estimates & model choice* (“A simulation analysis in a box” for your study) * Boettiger et al. 2011 in review Carl Boettiger, UC Davis Release of Constraint 25/37
  • 46. Returning to our opening example. . . Carl Boettiger, UC Davis Release of Constraint 26/37
  • 47. Returning to our opening example. . . Carl Boettiger, UC Davis Release of Constraint 26/37
  • 48. Can a key innovation release selective constraint? Differing trait diversification Differing strengths of stabilizing rates, or selection? Carl Boettiger, UC Davis Release of Constraint 27/37
  • 49. Differing Trait Diversification Rate Model Carl Boettiger, UC Davis Release of Constraint 28/37
  • 50. Differing Stabilizing Selection Strength Carl Boettiger, UC Davis Release of Constraint 29/37
  • 51. So which is the better model? Differing Diversification Log Likelihood: -94 Parameters: 4 Differing Selection Log Likelihood: -92 Parameters: 4 Carl Boettiger, UC Davis Release of Constraint 30/37
  • 52. So which is the better model? Differing Diversification Log Likelihood: -94 Parameters: 4 Differing Selection Log Likelihood: -92 Parameters: 4 Carl Boettiger, UC Davis Release of Constraint 30/37
  • 53. So which is the better model? Differing Diversification Log Likelihood: -94 Parameters: 4 Differing Selection Log Likelihood: -92 Parameters: 4 Not enough power for this comparison in this trait. Carl Boettiger, UC Davis Release of Constraint 30/37
  • 54. So which is the better model? Differing Diversification Log Likelihood: -94 Parameters: 4 Differing Selection Log Likelihood: -92 Parameters: 4 Not enough power for this comparison in this trait. . . . must try harder. . . Carl Boettiger, UC Davis Release of Constraint 30/37
  • 55. A slightly simpler model comparison Differing trait diversification Differing strengths of stabilizing rates, or selection? Carl Boettiger, UC Davis Release of Constraint 31/37
  • 56. Differing Stabilizing Selection Strength Differing Trait Diversification Rate Carl Boettiger, UC Davis Release of Constraint 32/37
  • 57. Comparison Carl Boettiger, UC Davis Release of Constraint 33/37
  • 58. If we prefer to be Bayesian: MCMC These are posteriors, not likelihood bootstraps. Requires priors, etc. But, looks pretty much the same. Carl Boettiger, UC Davis Release of Constraint 34/37
  • 59. Conclusions 1 A new phylogenetic method Carl Boettiger, UC Davis Release of Constraint 35/37
  • 60. Conclusions 1 A new phylogenetic method 2 Complex models are dangerous with inadequate power Carl Boettiger, UC Davis Release of Constraint 35/37
  • 61. Conclusions 1 A new phylogenetic method 2 Complex models are dangerous with inadequate power 3 Have methods to quantify power and uncertainty. Carl Boettiger, UC Davis Release of Constraint 35/37
  • 62. Conclusions 1 A new phylogenetic method 2 Complex models are dangerous with inadequate power 3 Have methods to quantify power and uncertainty. 4 Hobbit-sized swords: Smaller sub-models for smaller data Carl Boettiger, UC Davis Release of Constraint 35/37
  • 63. Acknowledgements Peter Wainwright Wainwright Lab Graham Coop Peter Ralph Funding: Download the development version now: https://github.com/cboettig/wrightscape Slides and more our on new Lab Blog: http://wainwrightlab.wordpress.com Carl Boettiger, UC Davis Release of Constraint 36/37
  • 64. An Addendum on Reproducible Research Slides on http://wainwrightlab.wordpress.com See meta-data, script, version of code, data, recent changes. . . Carl Boettiger, UC Davis Release of Constraint 37/37