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Bias in estimates of
palaeobiodiversity: beyond the
  marine macrofossil record

        Graeme T. Lloyd
Modern diversity is vast…
…but all stems from a single ancestor
How did life get so diverse?




  Last
Universal
Common
Ancestor
                 ?
How did life get so diverse?




  Last
Universal
Common
Ancestor
Extant vs. fossil record

                   Red – lineage through time plot based
                   on extant data only

                   Navy – lineage through time plot
                   based on fossil data




                                        Ezard et al. 2011
Sepkoski’s compendia
But can we trust the curves?




David Raup




                      Miller 2009, after Raup 1972
Raup’s two solutions


           Subsampling




           Modeling




                         Alroy 2010; Lloyd 2012
Common cause hypothesis
Collating the data –early curves




                               Peters 2005
Beyond the marine macrofossils…
Collating the data –early curves

     ?                      ?
 Open ocean   Shelf     Terrestrial
Dinosaurs data –early curves
Collating the



 Open ocean   Shelf     Terrestrial
Dinosaurs – data –early curves
Collating the common cause
         Non-marine     Haq SL
         surface area




          Packages        Miller SL




  DBCs

DBFs


                                      Butler et al. 2011
Dinosaurs – data –early curves
Collating the sampling bias

                    Dinosaur-bearing localities

                    Palaeoarea covered

                    Phylogenetic diversity

                    Taxonomic diversity

                    Dinosaur-bearing formations




                                     Benton et al. 2011
Dinosaurs - subsampling
Collating the data –early curves
                Raw + ghosts
        Raw




                      Subsampled




                                   Lloyd et al. 2008
Dinosaurs - modeling curves
Collating the data –early




                           Lloyd 2012
Coccolithophores–early curves
Collating the data



 Open ocean    Shelf     Terrestrial
Coccolithophores–early curves
Collating the data




                           Lloyd et al. 2012
Coccolithophores–early curves
Collating the data


            vs.
Coccolithophores–early curves
Collating the data - database




                      Yellow pins – ODP/DSDP




                  Green pins – Land-based localities
Coccolithophores–early curves
Collating the data – species




                           Lloyd et al. 2012
Coccolithophores–early curves
Collating the data – sampling




                           Lloyd et al. 2012
Coccolithophores–early curves
    Collating the data – correlation
  richness
             rho = 0.02

                              richness
                                         sampling

             sampling
rho = 0.40                rho = 0.48




                                               Lloyd et al. 2012
Coccolithophores–early curves
Collating the data – subsampling



 J   K   Pg+Ng    J   K   Pg+Ng




                           Lloyd et al. 2012
Coccolithophores–early curves
Collating the data – modeling



 J   K   Pg+Ng    J    K   Pg+Ng




                            Lloyd et al. 2012
Fishes the data –early curves
Collating



 Open ocean    Shelf     Terrestrial
Fishes – the data –early curves
Collating global curves

    Agassiz 1843         Carroll 1988




    Benton 1993         Sepkoski 2002




                       Lloyd and Friedman, accepted
Fishes – the data –early curves
Collating global curves

    Agassiz 1843                  Carroll 1988




                                      R^2 = 0.74
    Benton 1993                  Sepkoski 2002

                   R^2 = 0.91




                                Lloyd and Friedman, accepted
Fishes - database–early curves
Collating the data




                      Lloyd and Friedman, accepted
Fishes – the data –early curves
Collating raw diversity




                       Lloyd and Friedman, accepted
Fishes – the data –early curves
Collating proxies

    Formations




    Localities




    Grids


                           Lloyd and Friedman,
                           accepted
Fishes – the data –early curves
Collating correlations

                                    Formations
                    rho = 0.64
       rho = 0.71




   Localities                          Grids




                                 Lloyd and Friedman, accepted
Fishes – the data –early curves
Collating multivariate modeling




                       Lloyd and Friedman, accepted
Fishes – the data –early curves
Collating subsampling




                       Lloyd and Friedman, accepted
Fishes – the data –early curves
Collating modeling

    Formations




    Localities




    Grids

                           Lloyd and Friedman,
                           accepted
Fishes – the data –early curves
Collating cartilage vs. bone




                       Lloyd and Friedman, accepted
Fishes – the data –early curves
Collating cartilage vs. bone
              rho = 0.83   rho = 0.44

Formations




             rho = 0.53    rho = 0.20

Localities




             rho = 0.56    rho = 0.21

Grids



                           Lloyd and Friedman, accepted
Conclusions data –early curves
Collating the
                           ?



        Sampling bias                  Sampling bias



                        Common cause
Acknowledgements
  Collating the data –early curves


   Michael     Steve      Katie        Matt     Dave      Rachel
   Benton     Brusatte    Davis     Friedman    Hone     Jennings




Davide         Paul      Marcello   Manabu     Andrew   James       Jeremy
Pisani       Pearson      Ruta      Sakamot     Smith   Tarver       Young
                                       o
Papers, data, R code etc.
Collating the data –early curves



  http://www.graemetlloyd.com/

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Bias in estimates of palaeobiodiversity: beyond the marine macrofossil record