Are our estimates of palaeobiodiversity more apparent than real?
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Are our estimates of palaeobiodiversity more apparent than real?
Graeme T. Lloyd
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Modern diversity is vast…
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…but all stems from a single ancestor
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How did life get so diverse?
LastUniversalCommonAncestor
?
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How did life get so diverse?
LastUniversalCommonAncestor
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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
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Extant vs. fossil record
Near et al. 2012
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Earliest diversity curves
Agassiz 1843
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Earliest diversity curves
Philips 1860
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Early diversity curves
Valentine 1970
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But can we trust the curves?
Miller 2009, after Raup 1972
David Raup
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Raup’s two solutions
Subsampling
Modeling
Alroy 2010; Lloyd 2012
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Sepkoski’s compendia
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The consensus paper
Sepkoski et al. 1981
Median trace fossil assemblage richness
Species richness(Zoological record data)
David Raup
Median open-marinecommunity species richness
Dolf Seilacher
Richard Bambach
Genus diversityDavid Raup
Family diversityJack Sepkoski
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Collating the data –early curves2000s: a return to Raup’s solutions
Alroy et al. 2001 Peters and Foote 2001
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Collating the data –early curves2000s: a return to Raup’s solutions
Alroy et al. 2001 Peters and Foote 2001
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Collating the data –early curvesSubsampling
Alroy et al. 2008 Alroy 2010
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Collating the data –early curvesModeling
Smith and McGowan 2007
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Collating the data –early curvesCommon cause hypothesis
Peters 2005
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Collating the data –early curves(Common) causal arrows
Genera/Packages
Sea-level
O
C
S
Sr
Hannisdal and Peters 2011
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Collating the data –early curvesThe rest of the story…
Open ocean Shelf Terrestrial
? ?
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Collating the data –early curvesDinosaurs
Open ocean Shelf Terrestrial
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Collating the data –early curvesDinosaurs - sampling
Taxonomic diversity
Phylogenetic diversity
Dinosaur-bearing formations
Dinosaur-bearing localities
Palaeoarea covered
Benton et al. 2011
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Collating the data –early curvesDinosaurs - subsampling
Lloyd et al. 2008
Raw
Raw + ghosts
Subsampled
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Collating the data –early curvesDinosaurs - modeling
Lloyd 2012
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Collating the data –early curvesDinosaurs – common cause
Butler et al. 2011
Packages
Haq SL
Miller SL
Non-marinesurface area
DBFs
DBCs
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Collating the data –early curvesCoccolithophores
Open ocean Shelf Terrestrial
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Collating the data –early curvesCoccolithophores
Lloyd et al. 2012
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Collating the data –early curvesCoccolithophores
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Collating the data –early curvesCoccolithophores - database
Yellow pins – ODP/DSDP
Green pins – Land-based localities
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Collating the data –early curvesCoccolithophores – sp.:genus
Lloyd et al. 2012
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Collating the data –early curvesCoccolithophores – sp.:genus
Lloyd et al. 2012
Species:Genus
N Sites
N Authors
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Collating the data –early curvesCoccolithophores – sp.:genus
Lloyd et al. 2012
Rarefied with respect to N localities Rarefied with respect to N taxonomists
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Collating the data –early curvesCoccolithophores
vs.
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Collating the data –early curvesCoccolithophores – species
Lloyd et al. 2012
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Collating the data –early curvesCoccolithophores – sampling
Lloyd et al. 2012
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Collating the data –early curvesCoccolithophores – correlation
Lloyd et al. 2012
rho = 0.02
rho = 0.48rho = 0.40
richness
richness
sampling
sampling
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Collating the data –early curvesCoccolithophores – subsampling
Lloyd et al. 2012
J K Pg+Ng J K Pg+Ng
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Collating the data –early curvesCoccolithophores – modeling
Lloyd et al. 2012
J K Pg+Ng J K Pg+Ng
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Collating the data –early curvesFishes
Open ocean Shelf Terrestrial
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Collating the data –early curvesFishes – old curves
Agassiz 1843
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Collating the data –early curvesFishes – old curves
Thomson 1977
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Agassiz 1843 Carroll 1988
Benton 1993 Sepkoski 2002
Collating the data –early curvesFishes – global curves
Lloyd and Friedman, submitted
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Agassiz 1843 Carroll 1988
Benton 1993 Sepkoski 2002
Collating the data –early curvesFishes – global curves
Lloyd and Friedman, submitted
R^2 = 0.91
R^2 = 0.78
R^2
= 0.
74
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Agassiz 1843 Carroll 1988
Benton 1993 Sepkoski 2002
Collating the data –early curvesFishes – global curves
Lloyd and Friedman, submitted
R^2 = -0.21
R^2 = 0.25
R^2 = -0.15
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Collating the data –early curvesFishes - database
Lloyd and Friedman, submitted
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Collating the data –early curvesFishes – collector curve
Lloyd and Friedman, submitted
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Collating the data –early curvesFishes – grids sampled
1 2 3 4
0.5-degree net Plot localities Identify grids Count grids
Lloyd and Friedman, submitted
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Collating the data –early curvesFishes – raw diversity
Lloyd and Friedman, submitted
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Collating the data –early curvesFishes – proxies
Formations
Localities
Grids
Lloyd and Friedman,submitted
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Collating the data –early curvesFishes – correlations
Formations
Localities Grids
Lloyd and Friedman, submitted
rho = 0.64
rho = 0.59
rho
= 0.
71
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Collating the data –early curvesFishes – multivariate modeling
Lloyd and Friedman, submitted
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Collating the data –early curvesFishes – subsampling
Lloyd and Friedman, submitted
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Collating the data –early curvesFishes – modeling
Lloyd and Friedman,submitted
Formations
Localities
Grids
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Collating the data –early curvesFishes – cartilage vs. bone
Lloyd and Friedman, submitted
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Collating the data –early curvesFishes – cartilage vs. bone
Lloyd and Friedman, submitted
Formations
Localities
Grids
rho = 0.83
rho = 0.53
rho = 0.56
rho = 0.44
rho = 0.20
rho = 0.21
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Collating the data –early curvesConclusions
Sampling bias
Common cause
Sampling bias
?
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Collating the data –early curvesAcknowledgements
AndrewSmith
PaulPearson
JeremyYoung
MichaelBenton
SteveBrusatte
KatieDavis
DaveHone
RachelJennings
DavidePisani
MarcelloRuta
ManabuSakamot
o
JamesTarver
MattFriedman
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Collating the data –early curvesFishes – bone vs. cartilage
Lloyd and Friedman, submitted