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
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