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Anne Duputié Models in Evolutionary Ecology May 23rd, 2013 Modelling range shifts in dynamic environments How can evolution enter the stage? Anne Duputié CEFE, Montpellier

Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

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Page 1: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié CEFE, Montpellier

Page 2: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Parmesan et al. Nature 1999

Tircis

1940-69

1970-97

1915-39

- Migration

Responses to environmental changes (& lack thereof)

65% of 35 non migrating butterflies have shifted their range northwards in <100 y

Page 3: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Zhu et al. GCB 2012

Northward shift

Southward shift

Expansion

Contraction

- Migration (or not)

No

rth

ern

bo

un

dar

y ch

ange

(d

eg la

titu

de

)

Southern boundary change (deg latitude)

Responses to environmental changes (& lack thereof)

only 20% of 92 North American tree species show a northward range shift (59%: contraction)

Page 4: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Bradshaw & Holzapfel PNAS 2001

1940-69

1970-97

Latitude (corrected for altitude)

Wyeomyia smithii (photo S Gray)

- Migration (or not) - Adaptation

Cri

tica

l ph

oto

per

iod

(h

)

Responses to environmental changes (& lack thereof)

Evolution of critical photoperiod for entering into diapause (heritable trait, h²=15-70%) within 25 years

Page 5: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left

Responses to environmental changes (& lack thereof)

Drosphila birchii

Fragmented populations; no available genetic variance to respond to stress

Hoffmann et al Science 2003

Page 6: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Distribution of Chamaechrista fasciculata

Responses to environmental changes (& lack thereof)

Page 7: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

pre

coci

ty

number of leaves

Responses to environmental changes (& lack thereof)

Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Distribution of Chamaechrista fasciculata

Page 8: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Future optimum

climate shift by 2035

Responses to environmental changes (& lack thereof)

pre

coci

ty

number of leaves

Chamaechrista fasciculata

Distribution of Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Page 9: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Future optimum

migration climate shift by 2035

Responses to environmental changes (& lack thereof)

pre

coci

ty

number of leaves

Chamaechrista fasciculata

Distribution of Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Page 10: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Future optimum

migration

Adaptation (no migration)

climate shift by 2035

Responses to environmental changes (& lack thereof)

pre

coci

ty

number of leaves

Chamaechrista fasciculata

Distribution of Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Page 11: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Future optimum

climate shift by 2035

Responses to environmental changes (& lack thereof)

pre

coci

ty

number of leaves

Chamaechrista fasciculata

Distribution of Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Phenotype distribution

Page 12: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Future optimum

climate shift by 2035

Responses to environmental changes (& lack thereof)

pre

coci

ty

number of leaves

Chamaechrista fasciculata

Distribution of Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Phenotype distribution

Page 13: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Future optimum

climate shift by 2035

Responses to environmental changes (& lack thereof)

pre

coci

ty

number of leaves

Chamaechrista fasciculata

Distribution of Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Phenotype distribution

Page 14: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001

Future optimum

climate shift by 2035

Responses to environmental changes (& lack thereof)

pre

coci

ty

number of leaves

Chamaechrista fasciculata

Distribution of Chamaechrista fasciculata

- Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits

Phenotype distribution

Correlations among traits slow down evolution

Page 15: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Modelling species distributions

Observed occurrences

Observed/inferred density

Probability of occurrence

Habitat suitability

Page 16: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Modelling species distributions

Observed occurrences

Observed/inferred density

Probability of occurrence

Habitat suitability

Page 17: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Modelling species distributions

Example: Fagus sylvatica, European beech

Page 18: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Modelling species distributions

Example: Fagus sylvatica, European beech

Page 19: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Environment

Probability of occurrence

Tmax Tmin Prec GDD

Modelling species distributions

?

Page 20: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

“Ph

en

om

en

olo

gica

l”

Environment

Probability of occurrence

Tmax Tmin Prec GDD

Modelling species distributions P

(occ

urr

ence

)

environment

Page 21: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

“Ph

en

om

en

olo

gica

l”

“Pro

cess

-bas

ed”

Environment

Probability of occurrence

Traits: reaction norms

Growth/ Survival…

(Fitness)

Tmax Tmin Prec GDD

Modelling species distributions P

(occ

urr

ence

)

environment

Trai

t

environment

Page 22: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

“Ph

en

om

en

olo

gica

l”

“Pro

cess

-bas

ed”

Environment

Probability of occurrence

Traits: reaction norms

Growth/ Survival…

(Fitness)

“Co

nce

ptu

al” Traits:

realised vs optimum

Fitness

Tmax Tmin Prec GDD

Modelling species distributions P

(occ

urr

ence

)

environment

Trai

t

environment

Fitn

ess

Matching trait/optimum

Page 23: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

“Ph

en

om

en

olo

gica

l”

“Pro

cess

-bas

ed”

Environment

Probability of occurrence

Traits: reaction norms

Growth/ Survival…

(Fitness)

“Co

nce

ptu

al” Traits:

realised vs optimum

Fitness

Ease of calibration

Understanding

Tmax Tmin Prec GDD

Modelling species distributions P

(occ

urr

ence

)

environment

Trai

t

environment

Fitn

ess

Matching trait/optimum

Trait evolution

Page 24: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Genetic adaptation and distribution ranges

1. Constraints to adaptation ? a conceptual model of trait adaptation on a shifting gradient

2. Evolution of trait reaction norms a process-based model of tree distribution ranges

Page 25: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Factors limiting distribution ranges: Topography Biotic interactions Demography Adaptation Migration

1. Responses to environmental changes: existing conceptual models

Brown AmNat 1974

Grinnell Auk 1917

Mimura & Aitken JEB 2009

« Fundamental » niche

Realised niche

Svenning & Skov EcolLett 2007

Page 26: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Factors limiting distribution ranges: Topography Biotic interactions Demography Adaptation Migration

1. Responses to environmental changes: existing conceptual models

Brown AmNat 1974

Grinnell Auk 1917

Mimura & Aitken JEB 2009

« Fundamental » niche

Realised niche

Svenning & Skov EcolLett 2007

Page 27: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Breeder’s equation: R=h² S

Available genetic variance

Selection strength

1. Responses to environmental changes: existing conceptual models Fi

tnes

s

(in

trin

sic

gro

wth

rat

e r)

Mean trait z

Selection gradient

Model: - One species - Quantitative trait evolves

Page 28: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Fitn

ess

(i

ntr

insi

c gr

ow

th r

ate

r)

Mean trait z

Selection gradient

Model: - One species - Quantitative trait evolves - Environmental gradient

Fitness r

1. Responses to environmental changes: existing conceptual models

Page 29: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Model: - One species - Quantitative trait evolves - Environmental gradient - Variable population density

Fitness r

Space

Den

sity

Coupling demography/adaptation

1. Responses to environmental changes: existing conceptual models

Coupling demography/adaptation

Page 30: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Model: - One species - Quantitative trait evolves - Environmental gradient - Variable population density

Fitness r

Space

Den

sity

Coupling demography/adaptation

1. Responses to environmental changes: existing conceptual models

Page 31: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Some results of this type of models: - No spatial heterogeneity: Maximal speed of environmental change (Lynch & Lande 1993)

- Can be generalised to several traits (Gomulkiewicz & Houle AmNat 2009)

0 2

12

2 2

G Gc

S e S

V Vk k r

V N V

too little genetic variance or too weak selection

low fecundity

small population

Extinction if:

1. Responses to environmental changes: existing conceptual models

Page 32: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Some results of this type of models: - No spatial heterogeneity - Spatial heterogeneity only (Kirkpatrick & Barton AmNat 1997)

1. Responses to environmental changes: existing conceptual models

Adaptation depends on VG and migration

Mea

n t

rait

optimum

realised

Space

Page 33: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

1. Responses to environmental changes: existing conceptual models

Adaptation depends on VG and migration

Wider distribution for intermediate migration rates

Space

Mea

n t

rait

D

ensi

ty

optimum

realised

Space

Some results of this type of models: - No spatial heterogeneity - Spatial heterogeneity only (Kirkpatrick & Barton AmNat 1997)

Page 34: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Some results of this type of models: - No spatial heterogeneity - Spatial heterogeneity only - Spatial and temporal heterogeneity (Pease et al Ecology 1989)

1. Responses to environmental changes: existing conceptual models

Space

Mea

n t

rait

D

ensi

ty

optimum

realised

Space

Page 35: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Some results of this type of models: - No spatial heterogeneity - Spatial heterogeneity only - Spatial and temporal heterogeneity (Pease et al Ecology 1989)

1. Responses to environmental changes: existing conceptual models

Space

Mea

n t

rait

D

ensi

ty

optimum

realised

Space Clines move as the environment changes. If persisting, the species shifts its range at the speed of the environmental change, with a lag.

Page 36: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Some results of this type of models: - No spatial heterogeneity - Spatial heterogeneity only - Spatial and temporal heterogeneity

what about genetic constraints in heterogeneous environments?

1. Responses to environmental changes: existing conceptual models

Page 37: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

1. Genetic correlations and range shifts: model ingredients

- One species - Fitness depends on several traits under stabilizing selection: S

Tra

it 1

Trait 2

Adaptive landscape

S

Page 38: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

1. Genetic correlations and range shifts: model ingredients

- One species - Fitness depends on several traits under stabilizing selection: S - Genetic variance: G

Tra

it 1

Trait 2

Adaptive landscape

S

G

Page 39: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

1. Genetic correlations and range shifts: model ingredients

- One species - Fitness depends on several traits under stabilizing selection: S - Genetic variance: G - Environmental gradient, slope b

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Space

optimum 1

optimum 2

Trai

t m

ean

Page 40: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

- One species - Fitness depends on several traits under stabilizing selection: S - Genetic variance: G - Environmental gradient, slope b - Shifting at speed v

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Space

optimum 1

optimum 2

Trai

t m

ean

1. Genetic correlations and range shifts: model ingredients

Page 41: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

- One species - Fitness depends on several traits under stabilizing selection: S - Genetic variance: G - Environmental gradient, slope b - Shifting at speed v - Migration: density-dependent diffusion, σ

Tra

it 1

Trait 2

Adaptive landscape

S

G b σ

Space

Den

sity

Space

optimum 1

optimum 2

Trai

t m

ean

1. Genetic correlations and range shifts: model ingredients

Page 42: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

- One species - Fitness depends on several traits under stabilizing selection: S - Genetic variance: G - Environmental gradient, slope b - Shifting at speed v - Migration: density-dependent diffusion, σ - Spatial selection gradient Sb

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Space

optimum 1

optimum 2

Trai

t m

ean

Space

Den

sity

1. Genetic correlations and range shifts: model ingredients

Page 43: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

- One species - Fitness depends on several traits under stabilizing selection: S - Genetic variance: G - Environmental gradient, slope b - Shifting at speed v - Migration: density-dependent diffusion, σ - Spatial selection gradient Sb - G, S, b assumed constant

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Space

optimum 1

optimum 2

Trai

t m

ean

D

ensi

ty

Space

1. Genetic correlations and range shifts: model ingredients

Page 44: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Space

trait z optima

1. Genetic correlations and range shifts: results

Page 45: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Traits develop clines Clines often flatter than optima

Space

trait z optima

realised

1. Genetic correlations and range shifts: results

Page 46: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Traits develop clines Clines often flatter than optima Population density is gaussian

Space

trait z optima

fitness r

Space

Space

density n

realised

1. Genetic correlations and range shifts: results

Page 47: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Traits develop clines, shifting across time Clines often flatter than optima Population density is gaussian Population shifts

Space

trait z optima

fitness r

Space

Space

density n

realised

1. Genetic correlations and range shifts: results

Page 48: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Traits develop clines, shifting across time Clines often flatter than optima Population density is gaussian Population shifts, with constant lag

Space

trait z optima

fitness r

Space

Space

density n

realised

Ln

1. Genetic correlations and range shifts: results

Page 49: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Traits develop clines, shifting across time Clines often flatter than optima Population density is gaussian Population shifts, with constant lag

Space

trait z optima

fitness r

Space

Space

density n

realised

Ln ρ

1. Genetic correlations and range shifts: results

Page 50: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

σ

Traits develop clines, shifting across time Clines often flatter than optima Population density is gaussian Population shifts, with constant lag Range width constant

Space

trait z optima

fitness r

Space

Space

density n

realised

Ln ρ

Vn

1. Genetic correlations and range shifts: results

Page 51: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

G b

Sb

Traits develop clines, shifting across time Clines often flatter than optima Population density is gaussian Population shifts, with constant lag Range width constant

Analytical expressions for adaptation & demography. Increase when: Maximal adaptability A = bTS G Sb

G aligned with Sb Minimal spatial fitness gradient B = bT S b

b aligned with S

1. Genetic correlations and range shifts: results

Page 52: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Tra

it 1

Trait 2

Adaptive landscape

S

b

Traits develop clines, shifting across time Clines often flatter than optima Population density is gaussian Population shifts, with constant lag Range width constant

Analytical expressions for adaptation & demography. Increase when: Maximal adaptability A = bTS G Sb

G aligned with Sb Minimal spatial fitness gradient B = bT S b

b aligned with S

1. Genetic correlations and range shifts: results

Page 53: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

0 22 2c

B Av r

B

Extinction if change faster than:

Low fecundity Maladapted migrants

Not enough adaptation

Slow migration

Tolerance to change:

1. Genetic correlations and range shifts: results

Page 54: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

1. Genetic correlations and range shifts: results

Hinders adaptation Widens range

diffusion σ

Ran

ge w

idth

diffusion σ

Cri

tica

l sp

eed

o

f ch

ange

Migration:

diffusion σ

Clin

e sl

op

es

Maximal tolerance for intermediate dispersal

+ =

0 22 2c

B Av r

B

Extinction if change faster than:

Low fecundity Maladapted migrants

Not enough adaptation

Slow migration

Tolerance to change:

Page 55: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012

Adaptation to change easier when - genetic variance available in the direction of the (spatial) selection gradient - optimum changes in a direction under weak stabilising selection.

Counter gradients may appear due to genetic correlations/correlational selection

The more traits, the more persistence is threatened.

1. Genetic correlations and range shifts: wrap-up

Page 56: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Fixed genetic variance Fixed, diffusive dispersal Constrained fitness function No phenotypic plasticity Linear gradients shifting at constant speed

1. BUT…

Page 57: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Fixed genetic variance Fixed, diffusive dispersal Constrained fitness function No phenotypic plasticity Linear gradients shifting at constant speed

Burrows et al. Science 2011

Williams et al PNAS 2007

Non-analogous climates, B2 scenario

Projected temperature changes (°C/decade)

Spatial gradient (°C/km)

1. BUT…

Page 58: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Fixed genetic variance Fixed, diffusive dispersal Constrained fitness function No phenotypic plasticity Linear gradients shifting at constant speed

use a process-based model to: - evaluate selective pressures - take phenotypic plasticity into account - explicitly model spatial heterogeneity

1. BUT…

Page 59: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Genetic adaptation and distribution ranges

1. Constraints to adaptation ? a conceptual model of trait adaptation on a shifting gradient

2. Evolution of trait reaction norms a process-based model of tree distribution ranges

Page 60: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Fitness

Local climate

Phenological traits

Reproduction Survival

Resistance traits

Page 61: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Bud dormancy

Fitness

Reproduction Survival

Page 62: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Bud dormancy

Leafing

Flowering

Fitness

Reproduction Survival

Page 63: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Bud dormancy

Leafing

Flowering

Fruit maturation

Fitness

Reproduction Survival

Page 64: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Bud dormancy

Leafing

Flowering

Fruit maturation

Leaf senescence

Fitness

Reproduction Survival

Page 65: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Frost Bud dormancy

Leafing

Flowering

Fruit maturation

Leaf senescence

Fitness

Reproduction Survival

Page 66: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Frost

Drought Bud dormancy

Leafing

Flowering

Fruit maturation

Leaf senescence

Fitness

Reproduction Survival

Page 67: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001

2. Evolution of trait reaction norms: the model PHENOFIT

Frost

Drought Bud dormancy

Leafing

Flowering

Fruit maturation

Leaf senescence

Fitness

Reproduction Survival

Page 68: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: calibrating the model

fructification

leafing + senescence

Using time series: phenology & climate

Leaf

ing

dat

e observed

modelled

Year

Example: sessile oak Quercus petraea

Page 69: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: validating the model

Presence / absence Observed distribution Fitness simulated by

PHENOFIT (1980-2000)

Using observed distribution ranges

Example: sessile oak Quercus petraea

Page 70: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: model extrapolations

1950-2000 50-year fecundity, simulated by PHENOFIT

Under scenario A1Fi (“business as usual”)

Example: sessile oak Quercus petraea

Page 71: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

1990-2040

2. Evolution of trait reaction norms: model extrapolations

50-year fecundity, simulated by PHENOFIT

Under scenario A1Fi (“business as usual”)

Example: sessile oak Quercus petraea

Page 72: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2010-2060

2. Evolution of trait reaction norms: model extrapolations

50-year fecundity, simulated by PHENOFIT

Under scenario A1Fi (“business as usual”)

Example: sessile oak Quercus petraea

Page 73: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2030-2080

2. Evolution of trait reaction norms: model extrapolations

50-year fecundity, simulated by PHENOFIT

Under scenario A1Fi (“business as usual”)

Example: sessile oak Quercus petraea

Page 74: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2050-2100

2. Evolution of trait reaction norms: model extrapolations

50-year fecundity, simulated by PHENOFIT

Under scenario A1Fi (“business as usual”)

Example: sessile oak Quercus petraea

Page 75: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

« plastic » date

d=165

d=125

d=102

Method: impose event dates – e.g. leafing date.

Page 76: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

« plastic » date

d=165

d=125

d=102

d=166

d=126

d=103

d=164

d=124

d=101

1plasticd d 1plasticd d

Method: impose event dates – e.g. leafing date.

Page 77: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

« plastic » date

d=165

d=125

d=102

d=166

d=126

d=103

d=164

d=124

d=101

Fecundity

1plasticd d 1plasticd d

Method: impose event dates – e.g. leafing date.

Page 78: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

« plastic » date

d=165

d=125

d=102

d=166

d=126

d=103

d=164

d=124

d=101

Fecundity

1plasticd d 1plasticd d

log fecundity

trait

Method: impose event dates – e.g. leafing date.

Page 79: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

Sessile oak Quercus petraea

European beech Fagus sylvatica

Fecundity

Selection gradient

2000

Fecundity: high low

Budburst selected to occur: later earlier

Page 80: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2020

Sessile oak Quercus petraea

European beech Fagus sylvatica

Fecundity: high low

Budburst selected to occur: later earlier

Fecundity

Selection gradient

2. Evolution of trait reaction norms: determine selection gradients

Page 81: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

2040

Sessile oak Quercus petraea

European beech Fagus sylvatica

Fecundity

Selection gradient

Fecundity: high low

Budburst selected to occur: later earlier

Page 82: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

2060

Sessile oak Quercus petraea

European beech Fagus sylvatica

Fecundity

Selection gradient

Fecundity: high low

Budburst selected to occur: later earlier

Page 83: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

2080

Sessile oak Quercus petraea

European beech Fagus sylvatica

Fecundity

Selection gradient

Fecundity: high low

Budburst selected to occur: later earlier

Page 84: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

Sessile oak Quercus petraea

European beech Fagus sylvatica

Fecundity

Selection gradient

Fecundity: high low

Budburst selected to occur: later earlier

2100

Page 85: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

2. Evolution of trait reaction norms: determine selection gradients

Temperature

Pre

cip

itat

ion

s

Temperature

Selection for later budburst: western (warmer) part of the range

Sessile oak Quercus petraea

European beech Fagus sylvatica

Budburst selected to occur: later earlier

In the climatic (niche) space:

Page 86: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

Budburst date (imposed)

Fecu

nd

ity

Sessile oak Quercus petraea

Jan 30 Mar 30 Jun 20

2. Evolution of trait reaction norms: why these patterns?

Page 87: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

Budburst date (imposed)

Fecu

nd

ity

Sessile oak Quercus petraea

Jan 30 Mar 30 Jun 20

frost damage

2. Evolution of trait reaction norms: why these patterns?

Page 88: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep

insufficient time to reach maturation

Budburst date (imposed)

Fecu

nd

ity

Sessile oak Quercus petraea

Jan 30 Mar 30 Jun 20

frost damage

2. Evolution of trait reaction norms: why these patterns?

Page 89: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep

Method: suppress reaction norm phenology/local climate Treatments:

plastic population

d=165

d=125

d=102

d=152

d=120

d=96

year1 year 2

200

160

120

80

Resistance

Fitness

Climate

Phenology

2. Evolution of trait reaction norms: where is plasticity beneficial?

Page 90: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep

2. Evolution of trait reaction norms: where is plasticity beneficial?

Method: suppress reaction norm phenology/local climate Treatments:

plastic population no interannual plasticity

J=165

J=125

J=102

J=152

J=120

J=96

year1 year 2

200

160

120

80

d=145

d=125

d=102

d=145

d=125

d=102

Resistance

Fitness

Climate

Phenology

Page 91: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep

2. Evolution of trait reaction norms: where is plasticity beneficial?

advantageous burdensome

interannual plasticity

pre

cip

itat

ion

s

temperature

Page 92: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep

Pré

cip

itat

ion

s

Température

2. Evolution of trait reaction norms: where is plasticity beneficial?

advantageous burdensome

interannual plasticity

imposed budburst date

fecu

nd

ity

Page 93: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep

Pré

cip

itat

ion

s

Température

2. Evolution of trait reaction norms: where is plasticity beneficial?

advantageous burdensome

interannual plasticity

imposed budburst date

fecu

nd

ity

Page 94: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep

Pré

cip

itat

ion

s

Température

2. Evolution of trait reaction norms: where is plasticity beneficial?

advantageous burdensome

interannual plasticity

imposed budburst date

fecu

nd

ity

Page 95: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep

Pré

cip

itat

ion

s

Température

2. Evolution of trait reaction norms: where is plasticity beneficial?

advantageous burdensome

interannual plasticity

imposed budburst date

fecu

nd

ity

Page 96: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Wrap-up

Phenotypic plasticity may translate constraints Interannual variability on budburst/senescence dates weakly

impacts fitness + long-distance gene flow e.g. Kremer et al. 2012

-> reaction norms selected at the scale of the range?

… except at range/niche margins

e.g. Pichancourt & van Klinken 2012

Page 97: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Perspectives

Selection gradients vary over space/time weak response to climatic change?

Can the evolution of phenology mitigate projections of range shifts in temperate trees?

Optimal reaction norm

Simulated fitness, t=later

Page 98: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Perspectives

Selection gradients vary over space/time weak response to climatic change?

Can the evolution of phenology mitigate projections of range shifts in temperate trees?

Optimal reaction norm

Simulated fitness, t=later

PhD project, O. Ronce/I. Chuine, ED SIBAGHE

response Gβ

Page 99: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Perspectives

Selection gradients vary over space/time weak response to climatic change?

Can the evolution of phenology mitigate projections of range shifts in temperate trees?

Optimal reaction norm

realised reaction norm Simulated fitness, t=later

PhD project, O. Ronce/I. Chuine, ED SIBAGHE

response Gβ

Page 100: Anne Duputié | MEE 2013 | Modelling range shifts in dynamic environments – How can evolution enter the stage?

Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013

Thanks!

Isabelle Chuine

François Massol Ophélie Ronce

Alexis Rutschmann

Mark Kirkpatrick