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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 – 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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Zhu et al. GCB 2012
Northward shift
Southward shift
Expansion
Contraction
- Migration (or not)
Northernboundarychange(deglatitude)
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)
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
Criticalphotoperiod(h)
Responses to environmental changes (& lack thereof)
Evolution of critical photoperiod for
entering into diapause (heritable trait,
h²=15-70%) within 25 years
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
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)
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001
precocity
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
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)
precocity
number of leaves
Chamaechrista
fasciculata
Distribution of Chamaechrista fasciculata
- Migration (or not)
- Adaptation (or not)
. generation time
. no standing variance left
. correlations among traits
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001
Future optimum
migrationclimate shift
by 2035
Responses to environmental changes (& lack thereof)
precocity
number of leaves
Chamaechrista
fasciculata
Distribution of Chamaechrista fasciculata
- Migration (or not)
- Adaptation (or not)
. generation time
. no standing variance left
. correlations among traits
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)
precocity
number of leaves
Chamaechrista
fasciculata
Distribution of Chamaechrista fasciculata
- Migration (or not)
- Adaptation (or not)
. generation time
. no standing variance left
. correlations among traits
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)
precocity
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
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)
precocity
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
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)
precocity
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
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)
precocity
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Modelling species distributions
Observed occurrences
Observed/inferred density
Probability of occurrence
Habitat suitability
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Modelling species distributions
Observed occurrences
Observed/inferred density
Probability of occurrence
Habitat suitability
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Modelling species distributions
Example: Fagus sylvatica, European beech
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Modelling species distributions
Example: Fagus sylvatica, European beech
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Environment
Probability of occurrence
Tmax Tmin Prec GDD
…
Modelling species distributions
?
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
“Phenomenological”
Environment
Probability of occurrence
Tmax Tmin Prec GDD
…
Modelling species distributionsP(occurrence)
environment
“
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
“Phenomenological”
“Process-based”
Environment
Probability of occurrence
Traits:
reaction norms
Growth/
Survival…
(Fitness)
Tmax Tmin Prec GDD
…
Modelling species distributionsP(occurrence)
environment
Trait
environment
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
“Phenomenological”
“Process-based”
Environment
Probability of occurrence
Traits:
reaction norms
Growth/
Survival…
(Fitness)
“Conceptual”
Traits:
realised vs
optimum
Fitness
Tmax Tmin Prec GDD
…
Modelling species distributionsP(occurrence)
environment
Trait
environment
Fitness
Matching
trait/optimum
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
“Phenomenological”
“Process-based”
Environment
Probability of occurrence
Traits:
reaction norms
Growth/
Survival…
(Fitness)
“Conceptual”
Traits:
realised vs
optimum
Fitness
Ease of calibration
Understanding
Tmax Tmin Prec GDD
…
Modelling species distributionsP(occurrence)
environment
Trait
environment
Fitness
Matching
trait/optimum
Trait evolution
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
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
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
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
Fitness
(intrinsicgrowthrater)
Mean trait z
Selection
gradient
Model:
- One species
- Quantitative trait evolves
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Fitness
(intrinsicgrowthrater)
Mean trait z
Selection
gradient
Model:
- One species
- Quantitative trait evolves
- Environmental gradient
Fitness r
1. Responses to environmental changes: existing conceptual models
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Model:
- One species
- Quantitative trait evolves
- Environmental gradient
- Variable population density
Fitness r
Space
Density
Coupling demography/adaptation
1. Responses to environmental changes: existing conceptual models
Coupling demography/adaptation
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Model:
- One species
- Quantitative trait evolves
- Environmental gradient
- Variable population density
Fitness r
Space
Density
Coupling demography/adaptation
1. Responses to environmental changes: existing conceptual models
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
1
2
2 2
G G
c
S e S
V V
k 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
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
Meantrait
optimum
realised
Space
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
MeantraitDensity
optimum
realised
Space
Some results of this type of models:
- No spatial heterogeneity
- Spatial heterogeneity only (Kirkpatrick & Barton AmNat 1997)
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
MeantraitDensity
optimum
realised
Space
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
MeantraitDensity
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.
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
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
Trait1
Trait 2
Adaptive landscape
S
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
Trait1
Trait 2
Adaptive landscape
S
G
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
Trait1
Trait 2
Adaptive landscape
S
G
b
Space
optimum 1
optimum 2
Traitmean
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
Trait1
Trait 2
Adaptive landscape
S
G
b
Space
optimum 1
optimum 2
Traitmean
1. Genetic correlations and range shifts: model ingredients
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
Trait1
Trait 2
Adaptive landscape
S
G
bσ
Space
Density
Space
optimum 1
optimum 2
Traitmean
1. Genetic correlations and range shifts: model ingredients
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
Trait1
Trait 2
Adaptive landscape
S
G
b
Sb
σ
Space
optimum 1
optimum 2
Traitmean
Space
Density
1. Genetic correlations and range shifts: model ingredients
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
Trait1
Trait 2
Adaptive landscape
S
G
b
Sb
σ
Space
optimum 1
optimum 2
TraitmeanDensity
Space
1. Genetic correlations and range shifts: model ingredients
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
Trait 2
Adaptive landscape
S
G
b
Sb
σ
Space
trait z optima
1. Genetic correlations and range shifts: results
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
Trait1
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012
0 2
2 2
c
B A
v 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
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
Rangewidth
diffusion σ
Criticalspeed
ofchange
Migration:
diffusion σ
Clineslopes
Maximal tolerance for
intermediate dispersal
+ =
0 2
2 2
c
B A
v r
B



  Extinction if change faster than:
Low fecundity Maladapted
migrants
Not enough
adaptation
Slow migration
Tolerance to change:
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
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…
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…
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…
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
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
ReproductionSurvival
Resistance traits
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
ReproductionSurvival
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
ReproductionSurvival
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
ReproductionSurvival
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
ReproductionSurvival
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001
2. Evolution of trait reaction norms: the model PHENOFIT
FrostBud dormancy
Leafing
Flowering
Fruit maturation
Leaf
senescence
Fitness
ReproductionSurvival
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
ReproductionSurvival
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
ReproductionSurvival
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
Leafingdate
observed
modelled
Year
Example: sessile oak Quercus petraea
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
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
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
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
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
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
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.
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.
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.
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.
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
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
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
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
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
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep
2. Evolution of trait reaction norms: determine selection gradients
Temperature
Precipitations
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:
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep
Budburst date (imposed)
Fecundity
Sessile oak
Quercus petraea
Jan 30 Mar 30 Jun 20
2. Evolution of trait reaction norms: why these patterns?
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep
Budburst date (imposed)
Fecundity
Sessile oak
Quercus petraea
Jan 30 Mar 30 Jun 20
frost damage
2. Evolution of trait reaction norms: why these patterns?
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep
insufficient time to
reach maturation
Budburst date (imposed)
Fecundity
Sessile oak
Quercus petraea
Jan 30 Mar 30 Jun 20
frost damage
2. Evolution of trait reaction norms: why these patterns?
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?
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
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
precipitations
temperature
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep
Précipitations
Température
2. Evolution of trait reaction norms: where is plasticity beneficial?
advantageous
burdensome
interannual
plasticity
imposed budburst date
fecundity
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep
Précipitations
Température
2. Evolution of trait reaction norms: where is plasticity beneficial?
advantageous
burdensome
interannual
plasticity
imposed budburst date
fecundity
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep
Précipitations
Température
2. Evolution of trait reaction norms: where is plasticity beneficial?
advantageous
burdensome
interannual
plasticity
imposed budburst date
fecundity
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep
Précipitations
Température
2. Evolution of trait reaction norms: where is plasticity beneficial?
advantageous
burdensome
interannual
plasticity
imposed budburst date
fecundity
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
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
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
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
Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013
Thanks!
Isabelle Chuine
François Massol Ophélie Ronce
Alexis Rutschmann
Mark Kirkpatrick

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

  • 1. 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
  • 2. 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
  • 3. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Zhu et al. GCB 2012 Northward shift Southward shift Expansion Contraction - Migration (or not) Northernboundarychange(deglatitude) 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)
  • 4. 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 Criticalphotoperiod(h) Responses to environmental changes (& lack thereof) Evolution of critical photoperiod for entering into diapause (heritable trait, h²=15-70%) within 25 years
  • 5. 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
  • 6. 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)
  • 7. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001 precocity 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
  • 8. 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) precocity number of leaves Chamaechrista fasciculata Distribution of Chamaechrista fasciculata - Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits
  • 9. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Etterson & Shaw Science 2001 Future optimum migrationclimate shift by 2035 Responses to environmental changes (& lack thereof) precocity number of leaves Chamaechrista fasciculata Distribution of Chamaechrista fasciculata - Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits
  • 10. 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) precocity number of leaves Chamaechrista fasciculata Distribution of Chamaechrista fasciculata - Migration (or not) - Adaptation (or not) . generation time . no standing variance left . correlations among traits
  • 11. 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) precocity 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
  • 12. 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) precocity 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
  • 13. 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) precocity 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
  • 14. 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) precocity 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
  • 15. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Modelling species distributions Observed occurrences Observed/inferred density Probability of occurrence Habitat suitability
  • 16. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Modelling species distributions Observed occurrences Observed/inferred density Probability of occurrence Habitat suitability
  • 17. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Modelling species distributions Example: Fagus sylvatica, European beech
  • 18. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Modelling species distributions Example: Fagus sylvatica, European beech
  • 19. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Environment Probability of occurrence Tmax Tmin Prec GDD … Modelling species distributions ?
  • 20. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 “Phenomenological” Environment Probability of occurrence Tmax Tmin Prec GDD … Modelling species distributionsP(occurrence) environment “
  • 21. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 “Phenomenological” “Process-based” Environment Probability of occurrence Traits: reaction norms Growth/ Survival… (Fitness) Tmax Tmin Prec GDD … Modelling species distributionsP(occurrence) environment Trait environment
  • 22. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 “Phenomenological” “Process-based” Environment Probability of occurrence Traits: reaction norms Growth/ Survival… (Fitness) “Conceptual” Traits: realised vs optimum Fitness Tmax Tmin Prec GDD … Modelling species distributionsP(occurrence) environment Trait environment Fitness Matching trait/optimum
  • 23. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 “Phenomenological” “Process-based” Environment Probability of occurrence Traits: reaction norms Growth/ Survival… (Fitness) “Conceptual” Traits: realised vs optimum Fitness Ease of calibration Understanding Tmax Tmin Prec GDD … Modelling species distributionsP(occurrence) environment Trait environment Fitness Matching trait/optimum Trait evolution
  • 24. 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
  • 25. 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
  • 26. 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
  • 27. 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 Fitness (intrinsicgrowthrater) Mean trait z Selection gradient Model: - One species - Quantitative trait evolves
  • 28. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Fitness (intrinsicgrowthrater) Mean trait z Selection gradient Model: - One species - Quantitative trait evolves - Environmental gradient Fitness r 1. Responses to environmental changes: existing conceptual models
  • 29. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Model: - One species - Quantitative trait evolves - Environmental gradient - Variable population density Fitness r Space Density Coupling demography/adaptation 1. Responses to environmental changes: existing conceptual models Coupling demography/adaptation
  • 30. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Model: - One species - Quantitative trait evolves - Environmental gradient - Variable population density Fitness r Space Density Coupling demography/adaptation 1. Responses to environmental changes: existing conceptual models
  • 31. 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 1 2 2 2 G G c S e S V V k 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
  • 32. 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 Meantrait optimum realised Space
  • 33. 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 MeantraitDensity optimum realised Space Some results of this type of models: - No spatial heterogeneity - Spatial heterogeneity only (Kirkpatrick & Barton AmNat 1997)
  • 34. 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 MeantraitDensity optimum realised Space
  • 35. 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 MeantraitDensity 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.
  • 36. 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
  • 37. 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 Trait1 Trait 2 Adaptive landscape S
  • 38. 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 Trait1 Trait 2 Adaptive landscape S G
  • 39. 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 Trait1 Trait 2 Adaptive landscape S G b Space optimum 1 optimum 2 Traitmean
  • 40. 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 Trait1 Trait 2 Adaptive landscape S G b Space optimum 1 optimum 2 Traitmean 1. Genetic correlations and range shifts: model ingredients
  • 41. 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 Trait1 Trait 2 Adaptive landscape S G bσ Space Density Space optimum 1 optimum 2 Traitmean 1. Genetic correlations and range shifts: model ingredients
  • 42. 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 Trait1 Trait 2 Adaptive landscape S G b Sb σ Space optimum 1 optimum 2 Traitmean Space Density 1. Genetic correlations and range shifts: model ingredients
  • 43. 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 Trait1 Trait 2 Adaptive landscape S G b Sb σ Space optimum 1 optimum 2 TraitmeanDensity Space 1. Genetic correlations and range shifts: model ingredients
  • 44. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 Trait 2 Adaptive landscape S G b Sb σ Space trait z optima 1. Genetic correlations and range shifts: results
  • 45. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 46. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 47. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 48. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 49. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 50. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 51. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 52. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 Trait1 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
  • 53. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. EcolLett. 2012 0 2 2 2 c B A v 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
  • 54. 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 Rangewidth diffusion σ Criticalspeed ofchange Migration: diffusion σ Clineslopes Maximal tolerance for intermediate dispersal + = 0 2 2 2 c B A v r B      Extinction if change faster than: Low fecundity Maladapted migrants Not enough adaptation Slow migration Tolerance to change:
  • 55. 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
  • 56. 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…
  • 57. 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…
  • 58. 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…
  • 59. 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
  • 60. 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 ReproductionSurvival Resistance traits
  • 61. 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 ReproductionSurvival
  • 62. 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 ReproductionSurvival
  • 63. 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 ReproductionSurvival
  • 64. 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 ReproductionSurvival
  • 65. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Chuine & Beaubien EcolLett. 2001 2. Evolution of trait reaction norms: the model PHENOFIT FrostBud dormancy Leafing Flowering Fruit maturation Leaf senescence Fitness ReproductionSurvival
  • 66. 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 ReproductionSurvival
  • 67. 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 ReproductionSurvival
  • 68. 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 Leafingdate observed modelled Year Example: sessile oak Quercus petraea
  • 69. 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
  • 70. 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
  • 71. 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
  • 72. 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
  • 73. 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
  • 74. 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
  • 75. 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.
  • 76. 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.
  • 77. 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.
  • 78. 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.
  • 79. 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
  • 80. 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
  • 81. 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
  • 82. 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
  • 83. 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
  • 84. 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
  • 85. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep 2. Evolution of trait reaction norms: determine selection gradients Temperature Precipitations 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:
  • 86. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep Budburst date (imposed) Fecundity Sessile oak Quercus petraea Jan 30 Mar 30 Jun 20 2. Evolution of trait reaction norms: why these patterns?
  • 87. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep Budburst date (imposed) Fecundity Sessile oak Quercus petraea Jan 30 Mar 30 Jun 20 frost damage 2. Evolution of trait reaction norms: why these patterns?
  • 88. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Duputié et al. in prep insufficient time to reach maturation Budburst date (imposed) Fecundity Sessile oak Quercus petraea Jan 30 Mar 30 Jun 20 frost damage 2. Evolution of trait reaction norms: why these patterns?
  • 89. 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?
  • 90. 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
  • 91. 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 precipitations temperature
  • 92. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep Précipitations Température 2. Evolution of trait reaction norms: where is plasticity beneficial? advantageous burdensome interannual plasticity imposed budburst date fecundity
  • 93. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep Précipitations Température 2. Evolution of trait reaction norms: where is plasticity beneficial? advantageous burdensome interannual plasticity imposed budburst date fecundity
  • 94. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep Précipitations Température 2. Evolution of trait reaction norms: where is plasticity beneficial? advantageous burdensome interannual plasticity imposed budburst date fecundity
  • 95. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Rutschmann et al. in prep Précipitations Température 2. Evolution of trait reaction norms: where is plasticity beneficial? advantageous burdensome interannual plasticity imposed budburst date fecundity
  • 96. 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
  • 97. 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
  • 98. 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
  • 99. 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
  • 100. Anne Duputié – Models in Evolutionary Ecology – May 23rd, 2013 Thanks! Isabelle Chuine François Massol Ophélie Ronce Alexis Rutschmann Mark Kirkpatrick