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Phylogeny and uncertainty in
 analyses of life span




                                                                    Photo: bramblejungle/flickr
           Owen R. Jones* and Fernando Colchero
           Max Planck Institute for Demographic Research, Rostock
           *jones@demogr.mpg.de, website: owenjon.es


7th June 2012, EvoDemo Workshop, MPIDR, Germany
de Magalhaes & Costa 2009 J. Evol. Biol.




                                           Robinson 2005 BTO Research Report 407
Grey partridge (Perdix perdix)




 Fulmar (Fulmarus glacialis)
Data issues: sample size
                                                         30

                                                         25




                                Max. observed lifespan
•   Maximum observed life
                                                         20
    span increases with
    sample size                                          15


•   Species with small sample                            10

    sizes are problematic                                5

                                                         0

                                                              0   20   40      60    80   100

                                                                       Sample size
Data issues: truncation/censoring
Birth/hatching   Death
Data issues: truncation/censoring
Birth/hatching   Death




    Truncation
Data issues: truncation/censoring
Birth/hatching      Death




    Truncation


                 Censoring
Trait evolution

‣ Closely related species tend to share
  similar trait values by inheritance
  (phylogenetic signal)

‣ Traits can also be similar due to similar life
  style (convergent evolution)
Trait evolution


Correlation can be due to the influence of
the trait in question, or an inherited
characteristic.
Aim

• To develop and test a statistical modelling
  framework that accounts for these data
  issues while controlling for phylogeny
The data set
• British Trust for Ornithology has carried out
  mark-capture-recovery since 1933
• Maximum recorded life span for >200 species
• Clutch size, number of broods, body mass



                                 Robinson 2005 BTO Research Report 407
Bird illustrations: RSPB
Cuckoo (Cuculus canorus)
Phylogeny:Thomas, GH 2008 Proc. R. Soc. B
Bird illustrations: RSPB
Bird illustrations: RSPB
Bird illustrations: RSPB
Bird illustrations: RSPB
Phylogenetic signal measures the amount that
phylogeny influences trait (0 - 1).
Pagel’s Lambda ~ 0.73
Ordinary least squares regression


                  50



                  20
Life span (yrs)




                  10


                  5



                  2



                       5   50          500   5000   1    2             5         10     20

                                Weight (g)              Effort (clutch size * broods)
Ordinary least squares regression


                  50



                  20
Life span (yrs)




                  10


                  5



                  2



                       5    50          500   5000   1    2             5         10     20

                                 Weight (g)              Effort (clutch size * broods)


                           R2 = 0.26                          R2 = 0.27
Phylogenetic correction

                              Independent contrasts
                                                Assumes Lambda = 1
                  50



                  20
Life span (yrs)




                  10


                  5



                  2



                       5      50          500      5000   1       2             5         10     20

                                   Weight (g)                    Effort (clutch size * broods)


                           R2 = 0.26 to <0.01                 R2 = 0.27 to <0.01
Phylogenetic correction

                              Independent contrasts                                                                                        Optimised PGLS
                                                Assumes Lambda = 1                                                                                  Lambda = 0.73
                  50                                                                                                    50



                  20                                                                                                    20
Life span (yrs)




                                                                                                      Life span (yrs)
                  10                                                                                                    10


                  5                                                                                                     5



                  2                                                                                                     2



                       5      50          500      5000   1       2             5         10     20                          5    50          500   5000    1        2             5         10     20

                                   Weight (g)                    Effort (clutch size * broods)                                         Weight (g)                   Effort (clutch size * broods)


                           R2 = 0.26 to <0.01                 R2 = 0.27 to <0.01                                                 R2 = 0.26 to 0.06              R2 = 0.27 to 0.07
Phylogenetic correction

                              Independent contrasts                                                                                        Optimised PGLS
                                                Assumes Lambda = 1                                                                                  Lambda = 0.73
                  50                                                                                                    50



                  20                                                                                                    20
Life span (yrs)




                                                                                                      Life span (yrs)
                  10                                                                                                    10


                  5                                                                                                     5



                  2                                                                                                     2



                       5      50          500      5000   1       2             5         10     20                          5    50          500   5000    1        2             5         10     20

                                   Weight (g)                    Effort (clutch size * broods)                                         Weight (g)                   Effort (clutch size * broods)


                           R2 = 0.26 to <0.01                 R2 = 0.27 to <0.01                                                 R2 = 0.26 to 0.06              R2 = 0.27 to 0.07


                                     Can we improve the fit by accounting
                                             for data problems?
State-space model
Process model
 Predictor         Observed Response

   X                      Y
             Phylogeny
State-space model
  Process model
    Predictor         Observed Response

      X                      Y
                Phylogeny

Data model

 •Sample size
 •Censoring             True Response
 •Truncation                Y*
State-space model
Maximise likelihood of both
     Process model
                                              •       MCMC framework
        Predictor         Observed Response
                                              •       Simultaneously estimates:
           X                     Y                •    Coefficients of process model
                    Phylogeny                     •    Phylogenetic signal
                                                  •    True response
   Data model                                     •    Error in process model
                                                  •    Error in data model
      •Sample size                                •    -> Degree of censoring,
      •Censoring            True Response              truncation and sample size
      •Truncation               Y*                     effects.
State-space regression
                               models
                  50



                  20
Life span (yrs)




                  10


                  5



                  2



                       5    50          500   5000   1      2             5         10     20

                                 Weight (g)                Effort (clutch size * broods)


                       R2 = 0.06 to 0.10                 R2 = 0.07 to 0.12
BTO data underestimates lifespan for
          many species
                             1000
                             800
 % difference in life span

                             600
                             400
                             200
                             0




                                    0   5    10      15   20

                                            Effort
BTO data underestimates lifespan for
          many species
                             1000
                             800
 % difference in life span

                             600
                             400
                             200
                             0




                                    0   5    10      15   20

                                            Effort
Conclusions

• Life history patterns are moderated by
  phylogeny
• Method of correction is fundamentally
  important
• Data issues can be solved
• Further analyses are in the pipeline!
ComPADRe        ComADRe       DATLife      MaDDaBa      BiDDaBa

                 MPIDR                              CNRS

Life spans
              Life tables          Recapture histories
Projection matrices
                                      Integral projection models
                  Age structures
Acknowledgements
 MPIDR Germany - Dr. Fernando Colchero, Dr. Dalia Conde Ovando, Dr. Alex Scheuerlein,
 Dr. Roberto Salguero-Gómez, Julia Barthold, Dr. Annette Baudisch, Prof. James W. Vaupel
 CNRS, France - Profs. Jean-Dominique Lebreton, Jean-Michel Gaillard
 British Trust for Ornithology, Max Planck Society
PHYLOGENETIC SIGNAL AS A
          NUISANCE
                                                                                                            ●


•   Apparently strong relationships




                                        5
                                                                                              ●             ●
                                                                                         ●          ●            ●


    can be misleading.
                                                                                             ●               ●
                                                                                             ● ●
                                                                                    ●                       ●●
                                                                                               ●●
                                                                                               ●        ●
                                                                          ●●                   ●             ●




                                        4
                                                                                    ●●
                                                                             ●       ● ●● ● ●   ●●
                                                                                                 ●
                                                                                      ●   ●
                                                                            ● ● ●   ●   ● ●●     ●

    Driven by few independent
                                                                                ●
•                                                          ●              ●
                                                                          ●
                                                                             ● ●
                                                                             ● ●
                                                                             ● ●
                                                                              ●
                                                                                   ●●
                                                                                     ● ●

                                                                                      ●
                                                                                            ●
                                                                                              ●
                                                                                                   ●                 ●
                                                                                                                      ●




                                        3
    events.
                                                             ●                 ●
                                                                ● ●   ● ●    ●       ●
                                                      ●●                  ●    ● ●
                                                          ●           ●            ●
                                                         ●         ● ●●              ●
                                                       ●       ●         ● ●     ●
                                                    ● ● ● ●● ●                     ● ●
                                                    ●          ●   ●
                                                                       ●

                                        2
    Effectively overestimating
                                                ●

•                                                   ●
                                                            ● ● ●●
                                                                ●
                                                                           ●●
                                                            ●

    degrees of freedom - that’s why
                                                        ●                 ● ●
                                                                            ●
                                                        ●
                                                    ●                 ●
                                                        ●
                                        1




    it is sometimes called                                    ●




    ‘phylogenetic pseudocorrelation’.       1                     2             3                   4                     5
PHYLOGENETIC SIGNAL AS A
      NUISANCE




            5
            4
            3
            2
            1




                1   2   3   4   5
PHYLOGENETIC SIGNAL AS A
          NUISANCE
                                                                                                                                    ●




                                    3.5
                                                                                                           ●                         ●
                                                                                                                                ●
                                                                              ●                    ●
                                                                              ●
                                                                          ●                            ●




                                    3.0
                                                          ●                                    ●                            ●
                                                                  ●                                ●               ●●
                                                                           ●
                                                                          ●                            ●                    ●
                                                                                               ●                            ●
                                                                               ●       ●                                ●
                                                                                           ●           ●           ●




                                    2.5
                                                           ●●

    Apparently weak relationships
                                                   ●                                                           ●
•                                                 ● ● ●
                                                          ●●
                                                                 ●       ● ●● ●
                                                                                 ●
                                                                                   ●
                                                                                      ●●
                                                                                         ● ●
                                                                                               ●
                                                    ● ●          ● ●●          ● ●●      ●

    can be misleading.

                                    2.0
                                                             ●       ●●     ●●
                                                                                 ● ● ●        ●●
                                                                 ●          ●         ●
                                                            ●●                              ●
                                                             ●       ●                          ●
                                                                            ●     ●               ●
                                                  ●      ●                             ●●
                                                                                      ●
                                                                                      ●




                                    1.5
                                                                ●
                                                      ●       ●        ●●             ●
                                                  ●                                           ●

    Within clade effects can be
                                                      ● ●●               ●
•                                             ●   ●                          ●      ●     ●
                                                                  ●    ●            ●
                                                                  ●        ●                   ●
                                    1.0                    ●
                                                                    ●                 ●

    strong.
                                                      ●
                                                    ●                ●      ●     ●
                                    0.5

                                                          ●
                                                                          ●
                                                                      ●


                                          1                   2                        3                       4                         5
PHYLOGENETIC SIGNAL AS A
      NUISANCE




            3.5
            3.0
            2.5
            2.0
            1.5
            1.0
            0.5




                  1   2   3   4   5
Future work
                    • Model tempo and mode of evolution of
                        life span and reproductive effort.

                    Constrained to an optimum                           Random walk                                   Niche separation
              15




                                                              15




                                                                                                            15
              10




                                                              10




                                                                                                            10
              5




                                                              5




                                                                                                            5
Trait value




                                                Trait value




                                                                                              Trait value
              0




                                                              0




                                                                                                            0
              −5




                                                              −5




                                                                                                            −5
              −10




                                                              −10




                                                                                                            −10
              −15




                                                              −15




                    0    10    20    30   40                        0   10   20     30   40                 −15   0   10    20     30    40

                              Time                                           Time                                           Time

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Phylogeny and uncertainty in analyses of life span

  • 1. Phylogeny and uncertainty in analyses of life span Photo: bramblejungle/flickr Owen R. Jones* and Fernando Colchero Max Planck Institute for Demographic Research, Rostock *jones@demogr.mpg.de, website: owenjon.es 7th June 2012, EvoDemo Workshop, MPIDR, Germany
  • 2. de Magalhaes & Costa 2009 J. Evol. Biol. Robinson 2005 BTO Research Report 407
  • 3. Grey partridge (Perdix perdix) Fulmar (Fulmarus glacialis)
  • 4. Data issues: sample size 30 25 Max. observed lifespan • Maximum observed life 20 span increases with sample size 15 • Species with small sample 10 sizes are problematic 5 0 0 20 40 60 80 100 Sample size
  • 8.
  • 9. Trait evolution ‣ Closely related species tend to share similar trait values by inheritance (phylogenetic signal) ‣ Traits can also be similar due to similar life style (convergent evolution)
  • 10. Trait evolution Correlation can be due to the influence of the trait in question, or an inherited characteristic.
  • 11. Aim • To develop and test a statistical modelling framework that accounts for these data issues while controlling for phylogeny
  • 12. The data set • British Trust for Ornithology has carried out mark-capture-recovery since 1933 • Maximum recorded life span for >200 species • Clutch size, number of broods, body mass Robinson 2005 BTO Research Report 407
  • 15. Phylogeny:Thomas, GH 2008 Proc. R. Soc. B
  • 20. Phylogenetic signal measures the amount that phylogeny influences trait (0 - 1). Pagel’s Lambda ~ 0.73
  • 21. Ordinary least squares regression 50 20 Life span (yrs) 10 5 2 5 50 500 5000 1 2 5 10 20 Weight (g) Effort (clutch size * broods)
  • 22. Ordinary least squares regression 50 20 Life span (yrs) 10 5 2 5 50 500 5000 1 2 5 10 20 Weight (g) Effort (clutch size * broods) R2 = 0.26 R2 = 0.27
  • 23. Phylogenetic correction Independent contrasts Assumes Lambda = 1 50 20 Life span (yrs) 10 5 2 5 50 500 5000 1 2 5 10 20 Weight (g) Effort (clutch size * broods) R2 = 0.26 to <0.01 R2 = 0.27 to <0.01
  • 24. Phylogenetic correction Independent contrasts Optimised PGLS Assumes Lambda = 1 Lambda = 0.73 50 50 20 20 Life span (yrs) Life span (yrs) 10 10 5 5 2 2 5 50 500 5000 1 2 5 10 20 5 50 500 5000 1 2 5 10 20 Weight (g) Effort (clutch size * broods) Weight (g) Effort (clutch size * broods) R2 = 0.26 to <0.01 R2 = 0.27 to <0.01 R2 = 0.26 to 0.06 R2 = 0.27 to 0.07
  • 25. Phylogenetic correction Independent contrasts Optimised PGLS Assumes Lambda = 1 Lambda = 0.73 50 50 20 20 Life span (yrs) Life span (yrs) 10 10 5 5 2 2 5 50 500 5000 1 2 5 10 20 5 50 500 5000 1 2 5 10 20 Weight (g) Effort (clutch size * broods) Weight (g) Effort (clutch size * broods) R2 = 0.26 to <0.01 R2 = 0.27 to <0.01 R2 = 0.26 to 0.06 R2 = 0.27 to 0.07 Can we improve the fit by accounting for data problems?
  • 26. State-space model Process model Predictor Observed Response X Y Phylogeny
  • 27. State-space model Process model Predictor Observed Response X Y Phylogeny Data model •Sample size •Censoring True Response •Truncation Y*
  • 28. State-space model Maximise likelihood of both Process model • MCMC framework Predictor Observed Response • Simultaneously estimates: X Y • Coefficients of process model Phylogeny • Phylogenetic signal • True response Data model • Error in process model • Error in data model •Sample size • -> Degree of censoring, •Censoring True Response truncation and sample size •Truncation Y* effects.
  • 29. State-space regression models 50 20 Life span (yrs) 10 5 2 5 50 500 5000 1 2 5 10 20 Weight (g) Effort (clutch size * broods) R2 = 0.06 to 0.10 R2 = 0.07 to 0.12
  • 30. BTO data underestimates lifespan for many species 1000 800 % difference in life span 600 400 200 0 0 5 10 15 20 Effort
  • 31. BTO data underestimates lifespan for many species 1000 800 % difference in life span 600 400 200 0 0 5 10 15 20 Effort
  • 32. Conclusions • Life history patterns are moderated by phylogeny • Method of correction is fundamentally important • Data issues can be solved • Further analyses are in the pipeline!
  • 33.
  • 34. ComPADRe ComADRe DATLife MaDDaBa BiDDaBa MPIDR CNRS Life spans Life tables Recapture histories Projection matrices Integral projection models Age structures
  • 35. Acknowledgements MPIDR Germany - Dr. Fernando Colchero, Dr. Dalia Conde Ovando, Dr. Alex Scheuerlein, Dr. Roberto Salguero-Gómez, Julia Barthold, Dr. Annette Baudisch, Prof. James W. Vaupel CNRS, France - Profs. Jean-Dominique Lebreton, Jean-Michel Gaillard British Trust for Ornithology, Max Planck Society
  • 36. PHYLOGENETIC SIGNAL AS A NUISANCE ● • Apparently strong relationships 5 ● ● ● ● ● can be misleading. ● ● ● ● ● ●● ●● ● ● ●● ● ● 4 ●● ● ● ●● ● ● ●● ● ● ● ● ● ● ● ● ●● ● Driven by few independent ● • ● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● 3 events. ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ● 2 Effectively overestimating ● • ● ● ● ●● ● ●● ● degrees of freedom - that’s why ● ● ● ● ● ● ● ● 1 it is sometimes called ● ‘phylogenetic pseudocorrelation’. 1 2 3 4 5
  • 37. PHYLOGENETIC SIGNAL AS A NUISANCE 5 4 3 2 1 1 2 3 4 5
  • 38. PHYLOGENETIC SIGNAL AS A NUISANCE ● 3.5 ● ● ● ● ● ● ● ● 3.0 ● ● ● ● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● 2.5 ●● Apparently weak relationships ● ● • ● ● ● ●● ● ● ●● ● ● ● ●● ● ● ● ● ● ● ●● ● ●● ● can be misleading. 2.0 ● ●● ●● ● ● ● ●● ● ● ● ●● ● ● ● ● ● ● ● ● ● ●● ● ● 1.5 ● ● ● ●● ● ● ● Within clade effects can be ● ●● ● • ● ● ● ● ● ● ● ● ● ● ● 1.0 ● ● ● strong. ● ● ● ● ● 0.5 ● ● ● 1 2 3 4 5
  • 39. PHYLOGENETIC SIGNAL AS A NUISANCE 3.5 3.0 2.5 2.0 1.5 1.0 0.5 1 2 3 4 5
  • 40. Future work • Model tempo and mode of evolution of life span and reproductive effort. Constrained to an optimum Random walk Niche separation 15 15 15 10 10 10 5 5 5 Trait value Trait value Trait value 0 0 0 −5 −5 −5 −10 −10 −10 −15 −15 0 10 20 30 40 0 10 20 30 40 −15 0 10 20 30 40 Time Time Time