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