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400-million years of 'fishes': a survey of
sampling biases based on the UK record




             Graeme T. Lloyd
              Matt Friedman
Extant Vertebrate Diversity




                                      ‘fishes’
                                    (ca. 28,000)
species counts from Nelson (2006)
Historical ‘Fish’ Diversity




Agassiz 1833-1843
Historical ‘Fish’ Diversity




Benton 1998 (data from Fossil Record 2)
The Database




 
     1678 occurrences
 
     100 localities
 
     683 species
 
     437 genera
 
     70 geologic formations
Collector Curve
Genus richness
Fish-bearing geologic formations
Fish-bearing localities
Grid occupancy

      1              2             3                4




 0.5-degree net   localities   grids sampled   grids occupied
Grid occupancy
Correlations



           Correlation              Raw (rho)       G.D. (rho)
 Genus richness vs. Formations        0.66            0.48
  Genus richness vs. Localities       0.67            0.59
   Genus richness vs. Grids           0.75            0.66

                         p > 0.05        p < 0.05
Sampling correction - rarefaction
Sampling correction - formations
Sampling correction - localities
Sampling correction - grids
Clade comparison
Chondrichthyes     Osteichthyes
Genus richness
Chondrichthyes   Osteichthyes
Modeling - formations
Chondrichthyes                 Osteichthyes




 Raw (rho)   G.D. (rho)   Raw (rho)      G.D. (rho)
   0.87        0.65         0.70           0.53


                              p > 0.05         p < 0.05
Modeling - localities
Chondrichthyes                 Osteichthyes




 Raw (rho)   G.D. (rho)   Raw (rho)      G.D. (rho)
   0.94        0.87         0.69           0.37


                              p > 0.05         p < 0.05
Modeling – grid occupancy
Chondrichthyes                 Osteichthyes




 Raw (rho)   G.D. (rho)   Raw (rho)      G.D. (rho)
   0.93        0.83         0.72           0.57


                              p > 0.05         p < 0.05
Global curves
        Agassiz 1833/43 –   Benton 1993 –
        European            Global
        Species             families




      Carroll 1988 –        Sepkoski 2002 –
      Global                Marine
      genera                genera
Dataset correlations



                    UK          Agassiz1        Benton           Carroll    Sepkoski2
     UK                            0.39           0.76            0.67         0.68




                                                                                             Spearman
 Agassiz1           0.43                          -0.11          -0.26         0.18
  Benton            0.61          -0.15                           0.52         0.72
   Carroll          0.51          -0.20           0.78                         0.38
 Sepkoski2          0.65           0.26           0.90            0.71
                                               Pearson
                             p > 0.05                 p < 0.05        1
                                                                        European taxa only
                                                                      2
                                                                        marine taxa only
(Correlations based on generalized differences of log10 data)
Dataset correlations



                    UK          Agassiz1        Benton           Carroll    Sepkoski2
     UK                            0.39           0.76            0.67         0.68




                                                                                             Spearman
 Agassiz1           0.43                          -0.11          -0.26         0.18
  Benton            0.61          -0.15                           0.52         0.72
   Carroll          0.51          -0.20           0.78                         0.38
 Sepkoski2          0.65           0.26           0.90            0.71
                                               Pearson
                             p > 0.05                 p < 0.05        1
                                                                        European taxa only
                                                                      2
                                                                        marine taxa only
(Correlations based on generalized differences of log10 data)
Dataset correlations



                    UK          Agassiz1        Benton           Carroll    Sepkoski2
     UK                            0.39           0.76            0.67         0.68




                                                                                             Spearman
 Agassiz1           0.43                          -0.11          -0.26         0.18
  Benton            0.61          -0.15                           0.52         0.72
   Carroll          0.51          -0.20           0.78                         0.38
 Sepkoski2          0.65           0.26           0.90            0.71
                                               Pearson
                             p > 0.05                 p < 0.05        1
                                                                        European taxa only
                                                                      2
                                                                        marine taxa only
(Correlations based on generalized differences of log10 data)
Dataset correlations



                    UK          Agassiz1        Benton           Carroll    Sepkoski2
     UK                            0.39           0.76            0.67         0.68




                                                                                             Spearman
 Agassiz1           0.43                          -0.11          -0.26         0.18
  Benton            0.61          -0.15                           0.52         0.72
   Carroll          0.51          -0.20           0.78                         0.38
 Sepkoski2          0.65           0.26           0.90            0.71
                                               Pearson
                             p > 0.05                 p < 0.05        1
                                                                        European taxa only
                                                                      2
                                                                        marine taxa only
(Correlations based on generalized differences of log10 data)
Le conclusions
• First sampling corrected curves of fish
  richness:
Le conclusions
• First sampling corrected curves of fish
  richness:
  • Devonian diversity is far lower than expected…
Le conclusions
• First sampling corrected curves of fish
  richness:
  • Devonian diversity is far lower than expected…
  • …yet Paleogene rise survives…
Le conclusions
• First sampling corrected curves of fish
  richness:
  • Devonian diversity is far lower than expected…
  • …yet Paleogene rise survives…
  • …Cenozoic true “age of fishes”?
Le conclusions
• First sampling corrected curves of fish
  richness:
  • Devonian diversity is far lower than expected…
  • …yet Paleogene rise survives…
  • …Cenozoic true “age of fishes”?
• Chondricthyans are more correlated with
  sampling than osteichthyans
Le conclusions
• First sampling corrected curves of fish
  richness:
  • Devonian diversity is far lower than expected…
  • …yet Paleogene rise survives…
  • …Cenozoic true “age of fishes”?
• Chondricthyans are more correlated with
  sampling than osteichthyans
• UK record accounts for one half to two-thirds of
  global signal
Le conclusions
• First sampling corrected curves of fish
  richness:
  • Devonian diversity is far lower than expected…
  • …yet Paleogene rise survives…
  • …Cenozoic true “age of fishes”?
• Chondricthyans are more correlated with
  sampling than osteichthyans
• UK record accounts for one half to two-thirds of
  global signal
• Estimates of global fish richness have changed
  substantially in last 170 years

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400-million years of 'fishes': a survey of sampling biases based on the UK record

  • 1. 400-million years of 'fishes': a survey of sampling biases based on the UK record Graeme T. Lloyd Matt Friedman
  • 2. Extant Vertebrate Diversity ‘fishes’ (ca. 28,000) species counts from Nelson (2006)
  • 4. Historical ‘Fish’ Diversity Benton 1998 (data from Fossil Record 2)
  • 5. The Database  1678 occurrences  100 localities  683 species  437 genera  70 geologic formations
  • 10. Grid occupancy 1 2 3 4 0.5-degree net localities grids sampled grids occupied
  • 12. Correlations Correlation Raw (rho) G.D. (rho) Genus richness vs. Formations 0.66 0.48 Genus richness vs. Localities 0.67 0.59 Genus richness vs. Grids 0.75 0.66 p > 0.05 p < 0.05
  • 13. Sampling correction - rarefaction
  • 14. Sampling correction - formations
  • 15. Sampling correction - localities
  • 19. Modeling - formations Chondrichthyes Osteichthyes Raw (rho) G.D. (rho) Raw (rho) G.D. (rho) 0.87 0.65 0.70 0.53 p > 0.05 p < 0.05
  • 20. Modeling - localities Chondrichthyes Osteichthyes Raw (rho) G.D. (rho) Raw (rho) G.D. (rho) 0.94 0.87 0.69 0.37 p > 0.05 p < 0.05
  • 21. Modeling – grid occupancy Chondrichthyes Osteichthyes Raw (rho) G.D. (rho) Raw (rho) G.D. (rho) 0.93 0.83 0.72 0.57 p > 0.05 p < 0.05
  • 22. Global curves Agassiz 1833/43 – Benton 1993 – European Global Species families Carroll 1988 – Sepkoski 2002 – Global Marine genera genera
  • 23. Dataset correlations UK Agassiz1 Benton Carroll Sepkoski2 UK 0.39 0.76 0.67 0.68 Spearman Agassiz1 0.43 -0.11 -0.26 0.18 Benton 0.61 -0.15 0.52 0.72 Carroll 0.51 -0.20 0.78 0.38 Sepkoski2 0.65 0.26 0.90 0.71 Pearson p > 0.05 p < 0.05 1 European taxa only 2 marine taxa only (Correlations based on generalized differences of log10 data)
  • 24. Dataset correlations UK Agassiz1 Benton Carroll Sepkoski2 UK 0.39 0.76 0.67 0.68 Spearman Agassiz1 0.43 -0.11 -0.26 0.18 Benton 0.61 -0.15 0.52 0.72 Carroll 0.51 -0.20 0.78 0.38 Sepkoski2 0.65 0.26 0.90 0.71 Pearson p > 0.05 p < 0.05 1 European taxa only 2 marine taxa only (Correlations based on generalized differences of log10 data)
  • 25. Dataset correlations UK Agassiz1 Benton Carroll Sepkoski2 UK 0.39 0.76 0.67 0.68 Spearman Agassiz1 0.43 -0.11 -0.26 0.18 Benton 0.61 -0.15 0.52 0.72 Carroll 0.51 -0.20 0.78 0.38 Sepkoski2 0.65 0.26 0.90 0.71 Pearson p > 0.05 p < 0.05 1 European taxa only 2 marine taxa only (Correlations based on generalized differences of log10 data)
  • 26. Dataset correlations UK Agassiz1 Benton Carroll Sepkoski2 UK 0.39 0.76 0.67 0.68 Spearman Agassiz1 0.43 -0.11 -0.26 0.18 Benton 0.61 -0.15 0.52 0.72 Carroll 0.51 -0.20 0.78 0.38 Sepkoski2 0.65 0.26 0.90 0.71 Pearson p > 0.05 p < 0.05 1 European taxa only 2 marine taxa only (Correlations based on generalized differences of log10 data)
  • 27. Le conclusions • First sampling corrected curves of fish richness:
  • 28. Le conclusions • First sampling corrected curves of fish richness: • Devonian diversity is far lower than expected…
  • 29. Le conclusions • First sampling corrected curves of fish richness: • Devonian diversity is far lower than expected… • …yet Paleogene rise survives…
  • 30. Le conclusions • First sampling corrected curves of fish richness: • Devonian diversity is far lower than expected… • …yet Paleogene rise survives… • …Cenozoic true “age of fishes”?
  • 31. Le conclusions • First sampling corrected curves of fish richness: • Devonian diversity is far lower than expected… • …yet Paleogene rise survives… • …Cenozoic true “age of fishes”? • Chondricthyans are more correlated with sampling than osteichthyans
  • 32. Le conclusions • First sampling corrected curves of fish richness: • Devonian diversity is far lower than expected… • …yet Paleogene rise survives… • …Cenozoic true “age of fishes”? • Chondricthyans are more correlated with sampling than osteichthyans • UK record accounts for one half to two-thirds of global signal
  • 33. Le conclusions • First sampling corrected curves of fish richness: • Devonian diversity is far lower than expected… • …yet Paleogene rise survives… • …Cenozoic true “age of fishes”? • Chondricthyans are more correlated with sampling than osteichthyans • UK record accounts for one half to two-thirds of global signal • Estimates of global fish richness have changed substantially in last 170 years