Low rate of lineage diversification

High rates of lineage diversification

Ancestral trait innovation

Evolutionary dead ends (e.g. specialization hypothesis)

Key innovation hypothesis for diversity

Low rate of trait diversification

no special name? Non-adaptive radiation

High rate of trait diversification

Adaptive divergence? Local adaptation?

Adaptive radiation

Niche conservatism

Hodges 95Nectar spurs in Aquilegia

Hodges and Arnold 1995 Proc Roy Soc.

Hodges 97 table

Hodges and Arnold 1995 Proc Roy Soc.

zygomorphiclaterally symmetric

actinomorphicradially symmetric

Sargent 2004 Proc. Roy. Soc. London B.

D>0: 14D<0: 5

Maddison 2006 Evolution

Maddison et al. 2007 Evolution

Maddison et al. 2007 Evolution

Parameter estimation on

simulated trees, N=500 taxa

Mayrose et al. 2011 Science

Anolis ecomorphs

Losos 98

Losos et al. 1998 Science

Losos 98 - 2a

Losos et al. 1998 Science

Losos 98 - 2a

Losos et al. 1998 Science

Losos 98 - 2a

Losos et al. 1998 Science

Glor et al. 2003 Evolution

Harmon 03

Har

mon

et

al.

2003

Sci

ence

lineage diversity index = sum(obs – exp)positive value = early accumulation of lineages

Measuring niche conservatism - phylogenetic signal

K: Blomberg et al. (2003) Evolution; examples: Ackerly, PNAS in review

Blomberg’s K: measures degree of similarity among close relatives, relative to expectations based on Brownian motion

K<<1 K~1 K>>1

convergence brownian conserved

Har

mon

et

al.

2003

Sci

ence

mean subclade disparity/total disparityhigh values = high within group relative to among group variance = low phylo signal

Morphological disparity index = sum(obs-exp): positive values= deep clades span similar trait range, i.e. convergence across clades and low signal

Harmon 03-3

Harmon et al. 2003 Science

early diversification -> greater phylogenetic signal

Diversification of height in maples, Ceanothus and silverswords

~30 mya

~45 mya

rate = 0.014 felsens 0.10 felsens 0.79 felsens

height data: Ackerly, unpubl., Hickman (1993), Wagner (1999) phylogenies: Renner et al .(2008), Hardig et al. (2000), Baldwin & Sanderson (1998)

~5.2 mya

Are there differences among clades in trait diversification (= disparification) rates

O’Meara et al. 2006

Nested ML test:Does a 2 rate model provide a sufficiently better fit than a 1 rate model?

Martin and Wainright 2011

Martin and Wainright 2011

Martin and Wainright 2011

Martin and Wainright 2011

Quantifying rates of phenotypic evolution

Haldane (1949) Evolution; Gingerich (1983) Science

1 darwin = change by factor of e

million yrs

Rates of phenotypic diversification under Brownian motion

time

var(x)

1 felsen =

1 Var(loge(trait)

)

million yrs

Ackerly, PNAS 2009

Rates of phenotypic diversification (estimated for Brownian motion model)

Rate (felsens) Leaf sizeHeight

Acer

Aesculus

Arbutoideae

Ceanothus

lobelioids

silverswords

North temperateCaliforniaHawai’i

Acer

Aesculus

Arbutoideae

Ceanothus

lobelioids

silverswords

±1 s.e.

Ackerly, PNAS 2009

Becerra

Becerra 2005 PNAS

Becerra LTTLineages-through-time (LTT) plot

Becerra 2005 PNAS

Inga

Richardson et al. 2001 Science

Phylica

Richardson et al. 2001 Nature