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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?
LastUniversalCommonAncestor
?
How did life get so diverse?
LastUniversalCommonAncestor
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?
Miller 2009, after Raup 1972
David Raup
Raup’s two solutions
Subsampling
Modeling
Alroy 2010; Lloyd 2012
Sepkoski’s compendia
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
Collating the data –early curves2000s: a return to Raup’s solutions
Alroy et al. 2001 Peters and Foote 2001
Collating the data –early curves2000s: a return to Raup’s solutions
Alroy et al. 2001 Peters and Foote 2001
Collating the data –early curvesSubsampling
Alroy et al. 2008 Alroy 2010
Collating the data –early curvesModeling
Smith and McGowan 2007
Collating the data –early curvesCommon cause hypothesis
Peters 2005
Collating the data –early curves(Common) causal arrows
Genera/Packages
Sea-level
O
C
S
Sr
Hannisdal and Peters 2011
Collating the data –early curvesThe rest of the story…
Open ocean Shelf Terrestrial
? ?
Collating the data –early curvesDinosaurs
Open ocean Shelf Terrestrial
Collating the data –early curvesDinosaurs - sampling
Taxonomic diversity
Phylogenetic diversity
Dinosaur-bearing formations
Dinosaur-bearing localities
Palaeoarea covered
Benton et al. 2011
Collating the data –early curvesDinosaurs - subsampling
Lloyd et al. 2008
Raw
Raw + ghosts
Subsampled
Collating the data –early curvesDinosaurs - modeling
Lloyd 2012
Collating the data –early curvesDinosaurs – common cause
Butler et al. 2011
Packages
Haq SL
Miller SL
Non-marinesurface area
DBFs
DBCs
Collating the data –early curvesCoccolithophores
Open ocean Shelf Terrestrial
Collating the data –early curvesCoccolithophores
Lloyd et al. 2012
Collating the data –early curvesCoccolithophores
Collating the data –early curvesCoccolithophores - database
Yellow pins – ODP/DSDP
Green pins – Land-based localities
Collating the data –early curvesCoccolithophores – sp.:genus
Lloyd et al. 2012
Collating the data –early curvesCoccolithophores – sp.:genus
Lloyd et al. 2012
Species:Genus
N Sites
N Authors
Collating the data –early curvesCoccolithophores – sp.:genus
Lloyd et al. 2012
Rarefied with respect to N localities Rarefied with respect to N taxonomists
Collating the data –early curvesCoccolithophores
vs.
Collating the data –early curvesCoccolithophores – species
Lloyd et al. 2012
Collating the data –early curvesCoccolithophores – sampling
Lloyd et al. 2012
Collating the data –early curvesCoccolithophores – correlation
Lloyd et al. 2012
rho = 0.02
rho = 0.48rho = 0.40
richness
richness
sampling
sampling
Collating the data –early curvesCoccolithophores – subsampling
Lloyd et al. 2012
J K Pg+Ng J K Pg+Ng
Collating the data –early curvesCoccolithophores – modeling
Lloyd et al. 2012
J K Pg+Ng J K Pg+Ng
Collating the data –early curvesFishes
Open ocean Shelf Terrestrial
Collating the data –early curvesFishes – old curves
Agassiz 1843
Collating the data –early curvesFishes – old curves
Thomson 1977
Agassiz 1843 Carroll 1988
Benton 1993 Sepkoski 2002
Collating the data –early curvesFishes – global curves
Lloyd and Friedman, submitted
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
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
Collating the data –early curvesFishes - database
Lloyd and Friedman, submitted
Collating the data –early curvesFishes – collector curve
Lloyd and Friedman, submitted
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
Collating the data –early curvesFishes – raw diversity
Lloyd and Friedman, submitted
Collating the data –early curvesFishes – proxies
Formations
Localities
Grids
Lloyd and Friedman,submitted
Collating the data –early curvesFishes – correlations
Formations
Localities Grids
Lloyd and Friedman, submitted
rho = 0.64
rho = 0.59
rho
= 0.
71
Collating the data –early curvesFishes – multivariate modeling
Lloyd and Friedman, submitted
Collating the data –early curvesFishes – subsampling
Lloyd and Friedman, submitted
Collating the data –early curvesFishes – modeling
Lloyd and Friedman,submitted
Formations
Localities
Grids
Collating the data –early curvesFishes – cartilage vs. bone
Lloyd and Friedman, submitted
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
Collating the data –early curvesConclusions
Sampling bias
Common cause
Sampling bias
?
Collating the data –early curvesAcknowledgements
AndrewSmith
PaulPearson
JeremyYoung
MichaelBenton
SteveBrusatte
KatieDavis
DaveHone
RachelJennings
DavidePisani
MarcelloRuta
ManabuSakamot
o
JamesTarver
MattFriedman
Collating the data –early curvesFishes – bone vs. cartilage
Lloyd and Friedman, submitted
