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Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates @GraemeTLloyd Graeme T. Lloyd @Friedman_Lab Matt Friedman

Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

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Page 1: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-binning rates of continuous character evolution on a phylogeny: the origin of

amniotes, not tetrapods, is characterised by increased rates

@GraemeTLloyd Graeme T. Lloyd

@Friedman_LabMatt Friedman

Page 2: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Acknowledgements

Chris Organ Graham Slater

Page 3: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Questions

Page 4: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Questions

1. How can we measure rates of evolution in acontinuous character over multiple time bins?

Page 5: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Questions

2. How did the tempo of tetrapod mandibleevolution change over their early evolution?

1. How can we measure rates of evolution in acontinuous character over multiple time bins?

Page 6: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Questions

2. How did the tempo of tetrapod mandibleevolution change over their early evolution?

3. How does phylogenetic time-scalechoice affect patterns of tempo?

1. How can we measure rates of evolution in acontinuous character over multiple time bins?

Page 7: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

How fast?

Page 8: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate categories

Cloutier 1991; Lloyd et al. 2012; Lloyd submitted

Page 9: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate categories

Cloutier 1991; Lloyd et al. 2012; Lloyd submitted

Page 10: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate categories

Cloutier 1991; Lloyd et al. 2012; Lloyd submitted

Page 11: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate categories

Cloutier 1991; Lloyd et al. 2012; Lloyd submitted

Page 12: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate categories

Cloutier 1991; Lloyd et al. 2012; Lloyd submitted

Page 13: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate categories

Cloutier 1991; Lloyd et al. 2012; Lloyd submitted

Page 14: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate data types

Simpson 1944,1953; Westoll 1949

Page 15: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate data types

Simpson 1944,1953; Westoll 1949

Page 16: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate data types

Simpson 1944,1953; Westoll 1949

Page 17: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate data types

Simpson 1944,1953; Westoll 1949

Page 18: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Discrete data: disparity and tempo

Davis et al. 2012; Lloyd et al. 2012; Lloyd submitted

Page 19: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Discrete data: disparity and tempo

Davis et al. 2012; Lloyd et al. 2012; Lloyd submitted

Page 20: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Discrete data: disparity and tempo

Claddis

Davis et al. 2012; Lloyd et al. 2012; Lloyd submitted

Page 21: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Continuous data: mode dominates

Page 22: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Continuous data: mode dominates

Traitvalue

Time

Page 23: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Early Burst

Late BurstTrai

t

Time

Continuous data: mode dominates

Page 24: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate

Time

Continuous rates

Page 25: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Rate

Time

Continuous rates

Page 26: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Raia et al. 2013

Continuous rates

Page 27: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Harmon et al. 2008

New method: guts

Page 28: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

New method: description

Page 29: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

New method: description

Page 30: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

New method: description

Page 31: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

New method: description

Page 32: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

S1 S2 S31

New method: description

Page 33: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Anderson et al. 2013

Data

Page 34: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Anderson et al. 2013

Data

Page 35: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-scaling trees

Page 36: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Lee et al 2014

Time-scaling trees: simultaneous

Page 37: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-scaling trees: a posteriori

paleotree

strap

Page 38: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-scaling trees: a posteriori

+

paleotree

strap

Page 39: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-scaling trees: a posteriori

+ =

paleotreestrap

Page 40: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-scaling trees: a posteriori

Arbitrary

Branch-sharingRuta et al 2006; Brusatte et al 2008

Arbitrary addition(s)Derstler 1982

Minimum branch-lengthLaurin 2004

Page 41: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-scaling trees: a posteriori

cal3Bapst 2013

Hedman methodLloyd and Friedman in prep

NowakNowak et al 2013

Arbitrary Probabilistic

Branch-sharingRuta et al 2006; Brusatte et al 2008

Arbitrary addition(s)Derstler 1982

Minimum branch-lengthLaurin 2004

Page 42: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Time-scaling trees: a posteriori

Hedman methodLloyd and Friedman in prep

Arbitrary Probabilistic

Branch-sharingRuta et al 2006; Brusatte et al 2008

Page 43: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

τ7

τ8

τ6

τ5

Traditional approach first

Ruta et al. 2006; Brusatte et al. 2008

Branch-sharing (“equal”)

Page 44: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

τ7

τ8

τ6

τ5

root age

τ7

τ6

τ5

Traditional approach first Share time with preceding(non-zero length) branch

Ruta et al. 2006; Brusatte et al. 2008

τ8

Branch-sharing (“equal”)

Page 45: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

τ6

τ5

τ4

τ2

τ1

τ3

Hedman 2010; Lloyd and Friedman in prep

τ0

Probabilistic time-scaling

Page 46: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

τ0

τ6

τ5

τ4

τ2

τ1

τ3

Age (Ma)

τ0

Probabilistic time-scaling

Hedman 2010; Lloyd and Friedman in prep

Page 47: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

“Equal” time-tree

Page 48: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Probabilistic time-tree

Page 49: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

“Equal”timetree

Results

Page 50: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

“Equal”timetree

Results

Page 51: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

“Equal” timetreeProbablistic timetree

Results

Page 52: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Results with error

Page 53: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Results with error

Page 54: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

S1 S2

Two-tempo evolution

Page 55: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Slater 2013

BrownianOrnstein-Uhlenbeck

Two-mode evolution

Page 56: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Conclusions

Page 57: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Conclusions

Page 58: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Conclusions

Page 59: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Conclusions

Page 60: Time-binning rates of continuous character evolution on a phylogeny: the origin of amniotes, not tetrapods, is characterised by increased rates

Conclusions