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Are our estimates of
palaeobiodiversity more apparent
           than real?

         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
Extant vs. fossil record




                           Near et al. 2012
Earliest diversity curves




                            Agassiz 1843
Earliest diversity curves




                            Philips 1860
Early diversity curves




                         Valentine 1970
But can we trust the curves?




David Raup




                      Miller 2009, after Raup 1972
Raup’s two solutions


           Subsampling




           Modeling




                         Alroy 2010; Lloyd 2012
Sepkoski’s compendia
The consensus paper
          Median trace fossil       Dolf Seilacher
         assemblage richness


            Species richness             David Raup
     (Zoological record data)


     Median open-marine          Richard Bambach
 community species richness

                                         David Raup
              Genus diversity


                                    Jack Sepkoski
            Family diversity

Sepkoski et al. 1981
2000s: a the data –early solutions
Collating return to Raup’s curves




   Alroy et al. 2001   Peters and Foote 2001
2000s: a the data –early solutions
Collating return to Raup’s curves




   Alroy et al. 2001   Peters and Foote 2001
Subsampling
Collating the data –early curves




   Alroy et al. 2008     Alroy 2010
Modeling data –early curves
Collating the




                     Smith and McGowan 2007
Common cause hypothesis
Collating the data –early curves




                               Peters 2005
(Common) causal–early curves
Collating the data arrows
      Genera/
      Packages


      O


      C


      S


      Sr


      Sea-
      level
                      Hannisdal and Peters 2011
The rest thethe story… curves
Collating  of data –early


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



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




          Packages        Miller SL




  DBCs

DBFs


                                      Butler et al. 2011
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
Coccolithophores–early curves
Collating the data - database




                      Yellow pins – ODP/DSDP




                  Green pins – Land-based localities
Coccolithophores–early curves
Collating the data – sp.:genus




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


                Species:Genus
                                    N Sites




                                    N Authors




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

Rarefied with respect to N localities   Rarefied with respect to N taxonomists




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


            vs.
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 old curves




                              Agassiz 1843
Fishes – the data –early curves
Collating old curves




                             Thomson 1977
Fishes – the data –early curves
Collating global curves

    Agassiz 1843          Carroll 1988




    Benton 1993          Sepkoski 2002




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

    Agassiz 1843      R^2 = -0.21      Carroll 1988
        R^2 = -0.15




    Benton 1993                       Sepkoski 2002




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




                      Lloyd and Friedman, submitted
Fishes – the data –early curves
Collating collector curve




                       Lloyd and Friedman, submitted
Fishes – the data –early curves
    Collating grids sampled

      1                2                 3                      4




0.5-degree net   Plot localities   Identify grids          Count grids
                                               Lloyd and Friedman, submitted
Fishes – the data –early curves
Collating raw diversity




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

    Formations




    Localities




    Grids


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

                                     Formations
                    rho = 0.64
       rho = 0.71




   Localities                           Grids




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




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




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

    Formations




    Localities




    Grids

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




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




                       Lloyd and Friedman, submitted

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Are our estimates of palaeobiodiversity more apparent than real?