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Bias in estimates of palaeobiodiversity: beyond the marine macrofossil record
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
Sepkoski’s compendia
But can we trust the curves?
Miller 2009, after Raup 1972
David Raup
Raup’s two solutions
Subsampling
Modeling
Alroy 2010; Lloyd 2012
Collating the data –early curvesCommon cause hypothesis
Peters 2005
Collating the data –early curvesBeyond the marine macrofossils…
Open ocean Shelf Terrestrial
? ?
Collating the data –early curvesDinosaurs
Open ocean Shelf Terrestrial
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 curvesDinosaurs – sampling bias
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 curvesCoccolithophores
Open ocean Shelf Terrestrial
Collating the data –early curvesCoccolithophores
Lloyd et al. 2012
Collating the data –early curvesCoccolithophores
vs.
Collating the data –early curvesCoccolithophores - database
Yellow pins – ODP/DSDP
Green pins – Land-based localities
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
Agassiz 1843 Carroll 1988
Benton 1993 Sepkoski 2002
Collating the data –early curvesFishes – global curves
Lloyd and Friedman, accepted
Agassiz 1843 Carroll 1988
Benton 1993 Sepkoski 2002
Collating the data –early curvesFishes – global curves
R^2 = 0.91
R^2
= 0.
74
R^2 = 0.78
Lloyd and Friedman, accepted
Collating the data –early curvesFishes - database
Lloyd and Friedman, accepted
Collating the data –early curvesFishes – raw diversity
Lloyd and Friedman, accepted
Collating the data –early curvesFishes – proxies
Formations
Localities
Grids
Lloyd and Friedman,accepted
Collating the data –early curvesFishes – correlations
Formations
Localities Grids
rho = 0.64
rho = 0.59
rho
= 0.
71
Lloyd and Friedman, accepted
Collating the data –early curvesFishes – multivariate modeling
Lloyd and Friedman, accepted
Collating the data –early curvesFishes – subsampling
Lloyd and Friedman, accepted
Collating the data –early curvesFishes – modeling
Formations
Localities
Grids
Lloyd and Friedman,accepted
Collating the data –early curvesFishes – cartilage vs. bone
Lloyd and Friedman, accepted
Collating the data –early curvesFishes – cartilage vs. bone
Formations
Localities
Grids
rho = 0.83
rho = 0.53
rho = 0.56
rho = 0.44
rho = 0.20
rho = 0.21
Lloyd and Friedman, accepted
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 curvesPapers, data, R code etc.
http://www.graemetlloyd.com/