Are palaeobiodiversity patterns   more apparent than real?         Graeme T. Lloyd
Modern diversity is vast…
…but all stems from a single ancestor
…but all stems from a single ancestor
How did life get so diverse?  LastUniversalCommonAncestor                 ?
How did life get so diverse?  LastUniversalCommonAncestor
Extant vs. fossil record                   Red – lineage through time plot based                   on extant data only    ...
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 richn...
2000s: a the data –early solutionsCollating return to Raup’s curves   Alroy et al. 2001   Peters and Foote 2001
2000s: a the data –early solutionsCollating return to Raup’s curves   Alroy et al. 2001   Peters and Foote 2001
SubsamplingCollating the data –early curves   Alroy et al. 2008     Alroy 2010
Modeling data –early curvesCollating the                     Smith and McGowan 2007
Common cause hypothesisCollating the data –early curves                               Peters 2005
(Common) causal–early curvesCollating the data arrows      Genera/      Packages      O      C      S      Sr      Sea-   ...
But this the data –early curvesCollating is only part of the story      ?                         ?  Open ocean     Shelf ...
Dinosaurs data –early curvesCollating the Open ocean   Shelf     Terrestrial
Dinosaurs - sampling curvesCollating the data –early                  Dinosaur-bearing localities                  Palaeoa...
Dinosaurs - subsamplingCollating the data –early curves                Raw + ghosts        Raw                      Subsam...
Dinosaurs - modeling curvesCollating the data –early                           Lloyd 2012
Dinosaurs – data –early curvesCollating the common cause         Non-marine     Haq SL         surface area          Packa...
Coccolithophores–early curvesCollating the data Open ocean    Shelf     Terrestrial
Coccolithophores–early curvesCollating the data                           Lloyd et al. 2012
Coccolithophores–early curvesCollating the data
Coccolithophores–early curvesCollating the data - database                      Yellow pins – ODP/DSDP                  Gr...
Coccolithophores–early curvesCollating the data – sp.:genus                           Lloyd et al. 2012
Coccolithophores–early curvesCollating the data – sp.:genus                Species:Genus                                  ...
Coccolithophores–early curves Collating the data – sp.:genusRarefied with respect to N localities   Rarefied with respect ...
Coccolithophores–early curvesCollating the data            vs.
Coccolithophores–early curvesCollating the data – species                           Lloyd et al. 2012
Coccolithophores–early curvesCollating the data – sampling                           Lloyd et al. 2012
Coccolithophores–early curves    Collating the data – correlation  richness             rho = 0.02                        ...
Coccolithophores–early curvesCollating the data – subsampling J   K   Pg+Ng    J   K   Pg+Ng                           Llo...
Coccolithophores–early curvesCollating the data – modeling J   K   Pg+Ng    J    K   Pg+Ng                            Lloy...
Fishes the data –early curvesCollating Open ocean    Shelf     Terrestrial
Fishes – the data –early curvesCollating old curves                              Agassiz 1843
Fishes – the data –early curvesCollating old curves                             Thomson 1977
Fishes – the data –early curvesCollating global curves    Agassiz 1843          Carroll 1988    Benton 1993          Sepko...
Fishes – the data –early curvesCollating global curves    Agassiz 1843                   Carroll 1988                     ...
Fishes – the data –early curvesCollating global curves    Agassiz 1843      R^2 = -0.21      Carroll 1988        R^2 = -0....
Fishes - database–early curvesCollating the data                      Lloyd and Friedman, submitted
Fishes – the data –early curvesCollating collector curve                       Lloyd and Friedman, submitted
Fishes – the data –early curves    Collating grids sampled      1                2                 3                      ...
Fishes – the data –early curvesCollating raw diversity                       Lloyd and Friedman, submitted
Fishes – the data –early curvesCollating proxies    Formations    Localities    Grids                           Lloyd and ...
Fishes – the data –early curvesCollating correlations                                     Formations                    rh...
Fishes – the data –early curvesCollating modeling                       Lloyd and Friedman, submitted
Fishes – the data –early curvesCollating subsampling                       Lloyd and Friedman, submitted
Fishes – the data –early curvesCollating modeling    Formations    Localities    Grids                           Lloyd and...
Fishes – the data –early curvesCollating bone vs. cartilage                       Lloyd and Friedman, submitted
Fishes – the data –early curvesCollating bone vs. cartilage              rho = 0.83    rho = 0.44Formations             rh...
Conclusions data –early curvesCollating the                           ?        Sampling bias                  Sampling bia...
Acknowledgements  Collating the data –early curves   Michael     Steve      Katie        Matt     Dave      Rachel   Bento...
Fishes – the data –early curvesCollating bone vs. cartilage                       Lloyd and Friedman, submitted
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Are palaeobiodiversity patterns more apparent than real?

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Are palaeobiodiversity patterns more apparent than real?

  1. 1. Are palaeobiodiversity patterns more apparent than real? Graeme T. Lloyd
  2. 2. Modern diversity is vast…
  3. 3. …but all stems from a single ancestor
  4. 4. …but all stems from a single ancestor
  5. 5. How did life get so diverse? LastUniversalCommonAncestor ?
  6. 6. How did life get so diverse? LastUniversalCommonAncestor
  7. 7. 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
  8. 8. Earliest diversity curves Agassiz 1843
  9. 9. Earliest diversity curves Philips 1860
  10. 10. Early diversity curves Valentine 1970
  11. 11. But can we trust the curves?David Raup Miller 2009, after Raup 1972
  12. 12. Raup’s two solutions Subsampling Modeling Alroy 2010; Lloyd 2012
  13. 13. Sepkoski’s compendia
  14. 14. 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 diversitySepkoski et al. 1981
  15. 15. 2000s: a the data –early solutionsCollating return to Raup’s curves Alroy et al. 2001 Peters and Foote 2001
  16. 16. 2000s: a the data –early solutionsCollating return to Raup’s curves Alroy et al. 2001 Peters and Foote 2001
  17. 17. SubsamplingCollating the data –early curves Alroy et al. 2008 Alroy 2010
  18. 18. Modeling data –early curvesCollating the Smith and McGowan 2007
  19. 19. Common cause hypothesisCollating the data –early curves Peters 2005
  20. 20. (Common) causal–early curvesCollating the data arrows Genera/ Packages O C S Sr Sea- level Hannisdal and Peters 2011
  21. 21. But this the data –early curvesCollating is only part of the story ? ? Open ocean Shelf Terrestrial
  22. 22. Dinosaurs data –early curvesCollating the Open ocean Shelf Terrestrial
  23. 23. Dinosaurs - sampling curvesCollating the data –early Dinosaur-bearing localities Palaeoarea covered Phylogenetic diversity Taxonomic diversity Dinosaur-bearing formations Benton et al. 2011
  24. 24. Dinosaurs - subsamplingCollating the data –early curves Raw + ghosts Raw Subsampled Lloyd et al. 2008
  25. 25. Dinosaurs - modeling curvesCollating the data –early Lloyd 2012
  26. 26. Dinosaurs – data –early curvesCollating the common cause Non-marine Haq SL surface area Packages Miller SL DBCsDBFs Butler et al. 2011
  27. 27. Coccolithophores–early curvesCollating the data Open ocean Shelf Terrestrial
  28. 28. Coccolithophores–early curvesCollating the data Lloyd et al. 2012
  29. 29. Coccolithophores–early curvesCollating the data
  30. 30. Coccolithophores–early curvesCollating the data - database Yellow pins – ODP/DSDP Green pins – Land-based localities
  31. 31. Coccolithophores–early curvesCollating the data – sp.:genus Lloyd et al. 2012
  32. 32. Coccolithophores–early curvesCollating the data – sp.:genus Species:Genus N Sites N Authors Lloyd et al. 2012
  33. 33. Coccolithophores–early curves Collating the data – sp.:genusRarefied with respect to N localities Rarefied with respect to N taxonomists Lloyd et al. 2012
  34. 34. Coccolithophores–early curvesCollating the data vs.
  35. 35. Coccolithophores–early curvesCollating the data – species Lloyd et al. 2012
  36. 36. Coccolithophores–early curvesCollating the data – sampling Lloyd et al. 2012
  37. 37. Coccolithophores–early curves Collating the data – correlation richness rho = 0.02 richness sampling samplingrho = 0.40 rho = 0.48 Lloyd et al. 2012
  38. 38. Coccolithophores–early curvesCollating the data – subsampling J K Pg+Ng J K Pg+Ng Lloyd et al. 2012
  39. 39. Coccolithophores–early curvesCollating the data – modeling J K Pg+Ng J K Pg+Ng Lloyd et al. 2012
  40. 40. Fishes the data –early curvesCollating Open ocean Shelf Terrestrial
  41. 41. Fishes – the data –early curvesCollating old curves Agassiz 1843
  42. 42. Fishes – the data –early curvesCollating old curves Thomson 1977
  43. 43. Fishes – the data –early curvesCollating global curves Agassiz 1843 Carroll 1988 Benton 1993 Sepkoski 2002 Lloyd and Friedman, submitted
  44. 44. Fishes – the data –early curvesCollating global curves Agassiz 1843 Carroll 1988 R^2 = 0.74 Benton 1993 Sepkoski 2002 R^2 = 0.91 Lloyd and Friedman, submitted
  45. 45. Fishes – the data –early curvesCollating global curves Agassiz 1843 R^2 = -0.21 Carroll 1988 R^2 = -0.15 Benton 1993 Sepkoski 2002 Lloyd and Friedman, submitted
  46. 46. Fishes - database–early curvesCollating the data Lloyd and Friedman, submitted
  47. 47. Fishes – the data –early curvesCollating collector curve Lloyd and Friedman, submitted
  48. 48. Fishes – the data –early curves Collating grids sampled 1 2 3 40.5-degree net Plot localities Identify grids Count grids Lloyd and Friedman, submitted
  49. 49. Fishes – the data –early curvesCollating raw diversity Lloyd and Friedman, submitted
  50. 50. Fishes – the data –early curvesCollating proxies Formations Localities Grids Lloyd and Friedman, submitted
  51. 51. Fishes – the data –early curvesCollating correlations Formations rho = 0.64 rho = 0.71 Localities Grids Lloyd and Friedman, submitted
  52. 52. Fishes – the data –early curvesCollating modeling Lloyd and Friedman, submitted
  53. 53. Fishes – the data –early curvesCollating subsampling Lloyd and Friedman, submitted
  54. 54. Fishes – the data –early curvesCollating modeling Formations Localities Grids Lloyd and Friedman, submitted
  55. 55. Fishes – the data –early curvesCollating bone vs. cartilage Lloyd and Friedman, submitted
  56. 56. Fishes – the data –early curvesCollating bone vs. cartilage rho = 0.83 rho = 0.44Formations rho = 0.53 rho = 0.20Localities rho = 0.56 rho = 0.21Grids Lloyd and Friedman, submitted
  57. 57. Conclusions data –early curvesCollating the ? Sampling bias Sampling bias Common cause
  58. 58. Acknowledgements Collating the data –early curves Michael Steve Katie Matt Dave Rachel Benton Brusatte Davis Friedman Hone JenningsDavide Paul Marcello Manabu Andrew James JeremyPisani Pearson Ruta Sakamot Smith Tarver Young o
  59. 59. Fishes – the data –early curvesCollating bone vs. cartilage Lloyd and Friedman, submitted

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