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Is there a bias in deep sea
     diversity patterns?




       Graeme T. Lloyd
Collaborators




Andrew       Jeremy       Paul
 Smith       Young       Pearson
Talk Outline
• Introduction
• Deep sea diversity patterns:
  –   Deep sea diversity through time
  –   Deep sea rock record through time
  –   Correlations
  –   Modeling
• Species-per-genus patterns:
  –   Species-per-genus through time
  –   Deep sea rock record through time
  –   Authors through time
  –   Correlations
  –   Modeling
• Other diversity curves
Introduction
Fossil record is our only record of the
         diversification of life
Diversity correlates with rock record on
                 land…




   Peters and Foote 2001   Smith and McGowan 2007
…but what about the deep sea?
• Widely regarded as the best record we have:
  – Larger spatial extent than land
  – Abundant and widespread taxa
  – Near continuous sedimentation
  – Well studied (e.g. DSDP/ODP/IODP)
The database
• Study group is Coccolithophores
• Novel compilation from North Atlantic
• Compiled from 40 years ODP/DSDP
  data
• >50,000 occs from >15000 samples
• High temporal resolution (nanno zones)

Questions:
• What is the deep sea diversity pattern?
• What is the deep sea rock pattern?
• Are the two correlated to any degree?
Deep sea species richness
Deep sea rock record
Correlation tests

• First both time series were log-transformed
• Long term test:
  – Simple correlation
• Short term tests:
  – First differences (absolute)
  – Moving average differences (relative to long
    term trend)
• Degree (rho) and significance (p) of
  correlations determined using Spearman
  rank
Deep sea long-term correlation
Deep sea short-term correlation I
Deep sea short-term correlation II
Modeling approach
Modeling results

Real data
Predicted
Deep sea conclusions
Questions:
• What is the deep sea diversity pattern?
  – Linear rise with short-term fluctuations
• What is the deep sea rock pattern?
  – Exponential rise with short-term fluctuations
• Are the two correlated to any degree?
  – Yes, strongly and significantly, long-term and short-
    term
  – N sites is a good predictor of species richness
Species per genus patterns
Higher taxa as species proxies

•   Used since the earliest diversity curves…
•   …and continue to be (e.g. Alroy et al. 2008)
•   Originally pragmatic (less data required)
•   Then argued that species are inadequate
•   But, adequacy of higher taxa to represent
    species-level patterns is essentially untested
Taxonomic level affects pattern
Flessa and Jablonski 1985
              • Only explicit test of
                species-to-higher
                taxon ratio
              • Compared families
                to number of named
                species in Zoo.
                Record (Raup 1976)
              • Pattern of change
                differs; families
                become more
                speciose
Our database is superior!
• Species are standardised (synonyms)
• Species are assigned to genera
• Species are often widespread
• Species are long-ranging
• Species are comparatively stable
  taxonomically
• Question: how does the species-to-genus
  ratio change over time?
Species per genus
Number of sites
Number of authors
Long-term correlation
N sites
N authors
Short-term correlation
Species per genus
N sites
Short-term correlation
Species per genus
N authors
Correlations and conclusions
• Both number of sites and number of authors
  significantly correlate with species-per-genus
• Which is the best explanatory model?
  – Akaike weights = a combined model
  – Variance partitioning = a combined model

• Genera are not an accurate proxy for species
• Shift in species-per-genus ratio is best explained
  by a combination of sampling and worker bias, with
  limited room for a biological explanation
Other diversity curves
Other diversity curves
Other diversity curves

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Is there a bias in deep sea diversity patterns?

  • 1. Is there a bias in deep sea diversity patterns? Graeme T. Lloyd
  • 2. Collaborators Andrew Jeremy Paul Smith Young Pearson
  • 3. Talk Outline • Introduction • Deep sea diversity patterns: – Deep sea diversity through time – Deep sea rock record through time – Correlations – Modeling • Species-per-genus patterns: – Species-per-genus through time – Deep sea rock record through time – Authors through time – Correlations – Modeling • Other diversity curves
  • 5. Fossil record is our only record of the diversification of life
  • 6. Diversity correlates with rock record on land… Peters and Foote 2001 Smith and McGowan 2007
  • 7. …but what about the deep sea? • Widely regarded as the best record we have: – Larger spatial extent than land – Abundant and widespread taxa – Near continuous sedimentation – Well studied (e.g. DSDP/ODP/IODP)
  • 8. The database • Study group is Coccolithophores • Novel compilation from North Atlantic • Compiled from 40 years ODP/DSDP data • >50,000 occs from >15000 samples • High temporal resolution (nanno zones) Questions: • What is the deep sea diversity pattern? • What is the deep sea rock pattern? • Are the two correlated to any degree?
  • 9. Deep sea species richness
  • 10. Deep sea rock record
  • 11. Correlation tests • First both time series were log-transformed • Long term test: – Simple correlation • Short term tests: – First differences (absolute) – Moving average differences (relative to long term trend) • Degree (rho) and significance (p) of correlations determined using Spearman rank
  • 12. Deep sea long-term correlation
  • 13. Deep sea short-term correlation I
  • 14. Deep sea short-term correlation II
  • 17. Deep sea conclusions Questions: • What is the deep sea diversity pattern? – Linear rise with short-term fluctuations • What is the deep sea rock pattern? – Exponential rise with short-term fluctuations • Are the two correlated to any degree? – Yes, strongly and significantly, long-term and short- term – N sites is a good predictor of species richness
  • 18. Species per genus patterns
  • 19. Higher taxa as species proxies • Used since the earliest diversity curves… • …and continue to be (e.g. Alroy et al. 2008) • Originally pragmatic (less data required) • Then argued that species are inadequate • But, adequacy of higher taxa to represent species-level patterns is essentially untested
  • 21. Flessa and Jablonski 1985 • Only explicit test of species-to-higher taxon ratio • Compared families to number of named species in Zoo. Record (Raup 1976) • Pattern of change differs; families become more speciose
  • 22. Our database is superior! • Species are standardised (synonyms) • Species are assigned to genera • Species are often widespread • Species are long-ranging • Species are comparatively stable taxonomically • Question: how does the species-to-genus ratio change over time?
  • 29. Correlations and conclusions • Both number of sites and number of authors significantly correlate with species-per-genus • Which is the best explanatory model? – Akaike weights = a combined model – Variance partitioning = a combined model • Genera are not an accurate proxy for species • Shift in species-per-genus ratio is best explained by a combination of sampling and worker bias, with limited room for a biological explanation