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 95 Nectar spurs in Aquilegia Hodges and Arnold 1995 Proc Roy Soc.

Hodges 97 table Hodges and Arnold 1995 Proc Roy Soc.

zygomorphic laterally symmetric actinomorphic radially symmetric

Sargent 2004 Proc. Roy. Soc. London B. D>0: 14 D<0: 5

Maddison 2006 Evolution

Maddison et al Evolution

Parameter estimation on simulated trees, N=500 taxa

Mayrose et al Science

Anolis ecomorphs

Losos 98 Losos et al Science

Losos a Losos et al Science

Losos a Losos et al Science

Losos a Losos et al Science

Glor et al Evolution

Harmon 03 Harmon et al Science 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<<1K~1K>>1 convergencebrownianconserved

Harmon et al Science mean subclade disparity/total disparity high 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 Science early diversification -> greater phylogenetic signal

Harmon et al Assign proportional weighting of alternative models that best fit data

Diversification of height in maples, Ceanothus and silverswords ~30 mya ~45 mya rate = felsens0.10 felsens0.79 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 Nested ML test: Does a 2 rate model provide a sufficiently better fit than a 1 rate model?

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(log e (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 temperate California Hawai’i Acer Aesculus Arbutoideae Ceanothus lobelioids silverswords ±1 s.e. Ackerly, PNAS 2009

C. cuneatus Ceanothus in California C. oliganthus C. cordulatus C. fresnensis ©Dean Taylor 2005 high low sclerophylly Sierra pine forest Coastal and foothill chaparral  niche  niche

E D phylogenetic overdispersion = ‘  first’ coldhot :: :: DD EE

E D E D D E D coldhot :: :: phylogenetic clustering = ‘  first’

C. cuneatus C. oliganthus C. cordulatus C. fresnensis ©Dean Taylor 2005 Sierra pine forest Coastal and foothill chaparral Cerastes Euceanothus

How do  -niche traits diverge during allopatric speciation? E1->  1 E2->  2 11 22  div. coexistence 1) alternative  states favored in 2 environments, coincidentally promote coexistence in modern communities.

How do  -niche traits diverge during allopatric speciation? 22 E1->  1 E2->  2 11 22 continued  div. coexistence 2)  differences evolved in secondary contact of sister taxa, prior to diversification of two clades; modern coexistence reflects ancestral character displacement; early habitat divergence undetectable due to rapid rates of evolution. 11 secondary contact

Losos 98-2

Becerra Becerra 2005 PNAS

Becerra LTT Lineages-through-time (LTT) plot Becerra 2005 PNAS

Inga Richardson et al Science

Phylica Richardson et al Nature