Rates of evolution

Quantifying evolutionary rates

the darwin 1 darwin is a change in the character by a factor of e in 1 million years

Haldane 1949. Evolution 3: 51-56.

ln (x2) – ln (x1)d = t

Measuring evolutionary rates

the darwin the haldane

x2

h = g

sp

x1sp

( ( )) -

Sp= pooled standard deviation = SS1 + SS2

n1 + n2 - 2

g= number of generations separating the samplesSSi = sum of squared deviations for sample ini = sample size of sample i

Measuring evolutionary rates

the darwin the haldane specifying subscripts

dx(a:b,c) x = p (phenotypic) or g (genetic)

a= 1 (length), 2 (area) or 3 (volume)

b= log10(years over which the change was measured)

c = log10(years over which the rate is specified)

Measuring evolutionary rates

the darwin the haldane specifying subscripts

dp(1:0.95,6.0) phenotypic

length

log10(9); period of 9 years

log10(1000000); expressed over 1 million years

Measuring evolutionary rates

the darwin the haldane specifying subscripts

hx(b) x = p (phenotypic) or g (genetic)

b= log10(generations over which the change was measured)

Sources of information on evolutionary rates

artificial selection experiments

Rezende et al. 2006. J. Exp. Biol. 209: 115-127.

House mouse – Mus musculus

Rezende et al. 2006. J. Exp. Biol. 209: 115-127.

control

control

control

control

select

select

select

select

control

control

control

control

select

select

select

select

36 generations

control

select

select

Rezende et al. 2006. J. Exp. Biol. 209: 115-127.

control

control

control

control

select

select

select

select

control

control

control

control

select

select

select

select

36 generations

Distance run+137%Adult body mass -13.6%

Fat content -24.5%Food intake +8%

VO2max+13.2%

Rezende et al. 2006. J. Exp. Biol. 209: 115-127.

Sources of information on evolutionary rates

artificial selection experiments ‘natural’ experiments

allochronic and synchronic designs

Grant & Grant 1995. Evolution 49: 241-245.

Medium ground finch – Geospiza fortis

rain

fall

(cm

)

0

5

10

15

20

25

1970 1975 1980 1985

1976

Geospiza fortis

200

400

600

800

1000

1200

1400

1975 1976 1977 1978

popu

latio

n si

ze

0

2

4

6

8

10

12

1975 1976 1977 1978

seed

abu

ndan

ce (g

m-2

)

Jamaican fever plant - Tribulus cistoides

4

5

6

1975 1976 1977 1978

size

of a

vaila

ble

seed

s

Jamaican fever plant - Tribulus cistoides

Geospiza fortis

1981-1982

1976-1977

-0.5 0 0.5

weightwing length

beak lengthbeak depthbeak width

tarsus size

-0.5 0 0.5

weightwing length

beak lengthbeak depthbeak width

tarsus size

selection differential

rain

fall

(cm

)

0

5

10

15

20

25

1970 1975 1980 1985

1976 1983

Geospiza fortis

1981-1982

1976-1977

-0.5 0 0.5

1984-1985

weightwing length

beak lengthbeak depthbeak width

tarsus size

-0.5 0 0.5

weightwing length

beak lengthbeak depthbeak width

tarsus size

weightwing length

beak lengthbeak depthbeak width

tarsus size

selection differential

Geospiza fortis

1976-1977

1984-1985

weight

weight

r = +32250 darwin ≈ +0.709 haldanes

r = -11750 darwin ≈ -0.375 haldanes

Sources of information on evolutionary rates

artificial selection experiments ‘natural’ experiments

allochronic and synchronic designs

House sparrow – Passer domesticus

Johnston & Selander 1964. Science 144: 548-550.

Johnston & Selander 1964. Science 144: 548-550.

1900

1933

1914

Bermuda

Hawaii

18521962/3 (<111 generations)

Johnston & Selander 1964. Science 144: 548-550.

40

30

20

400 500 600 700

Death Valley

VancouverHonoluluMexico City

Refle

ctan

ce (%

)

Wavelength (mm)

Constantin Wilhelm Lambert Gloger (1803-1863)

Johnston & Selander 1964. Science 144: 548-550.

79

77

75

Win

g le

ngth

(mm

)83

81Edmonton

Death Valley

Lawrence

Johnston & Selander 1964. Science 144: 548-550.

Wei

ght (

g)

32

25

26

30 35 40 45 50 55 60

30

28

Isophane

Mosquitofish – Gambusia affinis

Stearns 1983. Am. Zool. 23: 65-76.

Stearns 1983. Am. Zool. 23: 65-76.

19051974 (140 generations)

Stearns 1983. Am. Zool. 23: 65-76.

Trait Stable Fluct %diff

Females Age at maturity (days) 82.5 88.3 7

Length at maturity (mm) 19.5 18.9 3

Growth rate 0.0066 0.0071 8

Dry weight offspring (mg) 1.00 1.12 12

Males Age at maturity (days) 72.6 84.0 16

Length at maturity (mm) 17.0 18.0 6

Growth rate 0.020 0.018 11

Blackcap – Sylvia atricapilla

Berthold et al. 1992. Nature 360: 668-670.

Berthold et al. 1992. Nature 360: 668-670.

1960s1990s (20 generations)

Guppy – Poecilia reticulata

Reznick et al. 1997. Science 275: 1934-1937.

Reznick et al. 1997. Science 275: 1934-1937.

11 years (18.1 generations)

waterfall

high predationcommunity

low predationcommunity

Crenicichla alta

Rivulus hartii

Reznick et al. 1997. Science 275: 1934-1937.

Trait Control Exp. %diff

Females Age at maturity (days) 85.6 93.5 9

Mass at maturity (mg) 162.3 189.2 17

Males Age at maturity (days) 48.6 58.2 20

Mass at maturity (mg) 67.5 76.1 13

Brown anole - Anolis sagrei

Losos et al. 1997. Nature 387: 70-73.

Losos et al. 1997. Nature 387: 70-73.

1977-811991 (13 generations)

Losos et al. 1997. Nature 387: 70-73.

5.0

2.0

2.21 2.24 2.25

3.0

Relative hindlimb length (mm)

4.0

2.262.22 2.23

Ln p

erch

dia

met

er (c

m) Staniel Cay

Flat periwinkle - Littorina obtusata

Seeley 1986. Proc. Natl. Acad. Sci. USA 83: 6897-6901.

Common shore crab - Carcinus maenas

Seeley 1986. Proc. Natl. Acad. Sci. USA 83: 6897-6901.

18711982

Appledore island, 1871 Appledore island, 1982

Seeley 1986. Proc. Natl. Acad. Sci. USA 83: 6897-6901.

Ln s

pire

hei

ght

1871

1982-84

1.6

-1

Ln shell width1.8 2.0 2.2

0

1

Holy grass - Anthoxanthum odoratum

Snaydon & Davies 1972. Evolution 26: 390-405.

Snaydon & Davies 1972. Evolution 26: 390-405.

Rothamsted Park Grass Experiment 1856-now

Holy grass - Anthoxanthum odoratum

Snaydon & Davies 1972. Evolution 26: 390-405.

18631960-70s

- plant height - disease resistance- tolerance of pH- tolerance of Al

2

1Transect section

3

4

5

6

2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

Phen

olog

ical

sco

re

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

pH=5.2 pH=3.9

Sources of information on evolutionary rates

artificial selection experiments ‘natural’ experiments

allochronic and synchronic designs divergence and evolution

200

150

100500

trai

t val

ue

time

A

B

evolution rateA = 1evolution rateB = 0divergence rate = 1

200

150

100500

trai

t val

ue

time

A

B

evolution rateA = 1evolution rateB = 1divergence rate = 0

200

150

100500

trai

t val

ue

time

A

B

evolution rateA = 1evolution rateB = -1divergence rate = 2

Sources of information on evolutionary rates

artificial selection experiments ‘natural’ experiments

allochronic and synchronic designs divergence and evolution phenotypic and genetic change

Sources of information on evolutionary rates

artificial selection experiments ‘natural’ experiments

allochronic and synchronic designs divergence and evolution phenotypic and genetic change intrapolation and extrapolation

200

150

100500

trai

t val

ue

time

actual rate = 1, -1inferred rate = 0

200

150

100500

trai

t val

ue

time

actual rate = 2, 0inferred rate = 1

200

150

100500

trai

t val

ue

time

actual rate = ?,?,?,…inferred rate = 1

200

150

100500

trai

t val

ue

time

actual rate = 1, 0inferred rate = 1

Sources of information on evolutionary rates

artificial selection experiments ‘natural’ experiments paleontological record

Dinosaurs in the Paleobiology Database (www.paleodb.org)

MacFadden 1992. Fossil horses. Cambridge Univ. Press

500

400

300

200

100

50 40 30 20 1060

Body

mas

s (k

g)

Million years ago

Hyracotherium

Mesohippus

Merychippus

Equus

Nannippus

MacFadden 1992. Fossil horses. Cambridge Univ. Press

M1

M2

M3M4

MacFadden 1992. Fossil horses. Cambridge Univ. Press

Species pair Dt (Myr) M1 (d) M2 (d) M3 (d) M4 (d)

Equus simplicidens – E. complicatus 2.0 0.000 -0.014 0.115 0.054

Merychippus isonesis – Pliohippus permix 2.5 0.072 0.101 0.185 0.180

Epihippus gracilis – Mesohippus bairdii 14 0.023 0.037 0.032 0.035

… 2-14 0.000-0.157

0.005-0.146

0.012-0.185

0.000-0.247

Threespine stickleback – Gasterosteus doryssus

Bell 2006. Paleobiology 32: 562-577.

0.6

Time (generations)

0.8

1.0

1.2

1.4

80000 600040002000

Mea

n nu

mbe

r of d

orsa

l spi

nes

Bell 2006. Paleobiology 32: 562-577.

Comparing evolutionary rates

faster evolution in shorter intervals

+ 0.27

time

0 1 2 3 4 5 6 7 8

char

acte

r

0

2

4

6

8

10

-0.60 +2.2

+0.25

evolutionary rate (darwin) time interval

n range geom. mean range geom. mean

Selection experiments 8 12000-200000 58700 1.5-10yr 3.7yr

Colonization 104 0-79700 370 70-300yr 170yr

Post-pleistocene mammals 46 0.11-32.0 3.7 1000-10000yr 8200yr

Fossil animals 363 0-26.2 0.08 8000yr-350Myr 3.8Myr

Fossil invertebrata 135 0-3.7 0.07 0.3-350Myr 7.9Myr

Fossil vertebrata 228 0-26.2 0.08 8000yr-98Myr 1.6Myr

All combined 521 0-200000 0.73 1.5yr-350Myr 0.2Myr

Gingerich 1983. Science 222: 159-161.

Comparing evolutionary rates

faster evolution in shorter intervals does natural selection suffice to explain fossil change?

evolutionary rate (darwin) time interval

n range geom. mean range geom. mean

Selection experiments 8 12000-200000 58700 1.5-10yr 3.7yr

Colonization 104 0-79700 370 70-300yr 170yr

Post-pleistocene mammals 46 0.11-32.0 3.7 1000-10000yr 8200yr

Fossil animals 363 0-26.2 0.08 8000yr-350Myr 3.8Myr

Fossil invertebrata 135 0-3.7 0.07 0.3-350Myr 7.9Myr

Fossil vertebrata 228 0-26.2 0.08 8000yr-98Myr 1.6Myr

All combined 521 0-200000 0.73 1.5yr-350Myr 0.2Myr

Gingerich 1983. Science 222: 159-161.

Comparing evolutionary rates

faster evolution in shorter intervals does natural selection suffice to explain fossil change? does natural selection suffice to explain radiation?

Geospiza fortis

1976-1977: beak depth +5%

Geospiza magnirostris Geospiza fortis

23 bouts

volcanic island chain

crust

man

tle p

lum

e

asthenosphere

mid-ocean ridge

lithosphere

4 Mya

1-0.5 Mya

570 000 ya

Punctuated equilibrium vs phyletic gradualism

Eldredge & Gould’s original idea

character

time

character

phyletic gradualism punctuated equilibrium

Punctuated equilibrium vs phyletic gradualism

Eldredge & Gould’s original idea

character

punctuated equilibrium

stasis

stabilizing selection

constraintlack of genetic variation

lack of expressedgenetic variation

developmentalcanalization

Punctuated equilibrium vs phyletic gradualism

genotype 1

genotype 2

other extremeenvironment extreme

Phen

otyp

e

Punctuated equilibrium vs phyletic gradualism

Eldredge & Gould’s original idea Gradualism and constant rate

Punctuated equilibrium vs phyletic gradualism

Eldredge & Gould’s original idea Gradualism and constant rate Valley crossing: macromutations and hopefull monsters

character

punctuated equilibrium

revolution

allopatric speciation

genetic revolution

Evening primrose - Oenothera lamarckianaHugo de Vries (1848-1935)

Richard Goldschmidt (1878-1958)

Lymantria

wild type antennapedia proboscipedia/ antennapedia

Punctuated equilibrium vs phyletic gradualism

Eldredge & Gould’s original idea Gradualism and constant rate Valley crossing: macromutations and hopefull monsters Examining the facts

Paleontological evidence

Cheetham 1986. Paleobiology 12: 199-202.

Bryozoa

Cheetham 1986. Paleobiology 12: 199-202.

M. auriculatum M. colligatum M. lacrymosum M. new species

Metrarabdotos

Bryozoa (Metrarabdotus) of the Miocene-Pliocene period(100 populations; 1000 specimens; 46 characteristics)

Cheetham 1986. Paleobiology 12: 199-202.

morphological differentiation

Williamson 1981. Nature 293: 437-443.

modern lake

Mutela

Cenozoic molluscs of the Turkana basin(13 species, 3300 specimens)

CaelaturaBellamya

Williamson 1981. Nature 293: 437-443.

modern lake

freshwater deposits

Williamson 1981. Nature 293: 437-443.

modern lake

freshwater deposits

Van Bocxlaer et al. 2007. Evolution 62: 511-520

freshwater deposits

arid

relatively wetmarkedly wet

Sheldon 1987. Nature 330: 561-563.

Ordovician trilobites from central Wales(~15000 specimens)

Sheldon 1987. Nature 330: 561-563.

number of pygidial ribs

time

Punctuated equilibrium vs phyletic gradualism

Eldredge & Gould’s original idea Gradualism and constant rate Valley crossing: macromutations and hopefull monsters Examining the facts

Paleontological evidence Neontological evidence

A B C D

gradual evolution A and B are more similar than C and D

punctuated equilibrium A and B are as similar as C and D

gradual evolution genetic variation similar in rich and poor clades

punctuated equilibrium genetic variation greater in rich clades

gradual evolution model with unit branch lenghts fits worse

punctuated equilibrium model with unit brach lenghts fits better

Moeers et al. 1999. Am. Nat. 154: 249-259.

Moeers et al. 1999. Am. Nat. 154: 249-259.

plumage colorationunison calls

sternum and keel anatomy

Moeers et al. 1999. Am. Nat. 154: 249-259.

Grus japonensis

Grus americana

Grus grus

Grus monachus

Grus nigricollis

Grus leucogeranus

Grus antigone

Grus vipio

Grus rubicunda

Grus canadensis

Bugeranus carunculatus

Anthropoides paradisea

Anthropoides virgo

Moeers et al. 1999. Am. Nat. 154: 249-259.

trait model performance

plumage color gradual = speciational < non-historical

unison call gradual > speciational > non-historical

anatomy gradual = speciational > non-historical

Whittall & Hodges 2007. Nature 477: 706-710.

Comet orchid - Angraecum sesquipedale

…

P.S. I shall be very glad indeed to see an Arethusa & then I must stop. I have just received such a Box full from Mr Bateman with the astounding Angræcum sesquipedalia with a nectary a foot long— Good Heavens what insect can suck it— I will write about Dimorphism.

C.R. Darwin to J.D. Hooker, 25 January 1862

Whittall et al. 2007. Nature 477: 706-710.

Xanthopan morgani ssp. praedicta

Whittall et al. 2007. Nature 477: 706-710.

spur and tongue length

fitne

ss

spur length

time

Whittall et al. 2007. Nature 477: 706-710.

spur and tongue length

fitne

ss

spur length

time

Whittall et al. 2007. Nature 477: 706-710.

Whittall et al. 2007. Nature 477: 706-710.

BM-model: AIC=51.933PEAKSmodel: AIC=26.90

gradual < punctuated