Natural Selection and The Co-Evolution of Interactions

Natural selection and the
co-evolution of interactions
Structure:
• Coevolution and natural selection mediated by species
interactions
• The essential components of evolution through natural
selection
• Natural selection analysis
• Heritability and Evolutionary responses
• Trade-offs
• Agents of selection
• Local adaptation
• Geographic mosaic theory of coevolution
Structure:
interactions
selection
• Trade-offs
Species interactions
and co-evolution
• Species interactions influence the interacting organisms in two
different ways, ecologically and evolutionary.
• Both take place at the level of individuals
• Ecological effects on mean absolute fitness of individuals leads to

changes in numbers, and thus influence population dynamics,
abundances, distributions, and community composition
• Effects on the covariance between fitness and traits is the basis for
natural selection and evolution, i.e. changes in trait distributions
• Interactions can have an evolutionary but not an ecological effect,

or vice versa
Natural selection mediated by
species interactions
• Evolution takes place on an ecological stage; ecological interactions
between species and the biotic and abiotic aspects of their environment
are the drivers of adaptive evolution.
• Natural selection mediated by species interactions and co-evolution is

important, because in most environments species interactions have
stronger effects than abiotic factors.
• Most species compete with similar species
• Most species interact with other species as consumers (e.g. predators,

herbivores or parasites), food resources, or both.
• Many species are engaged in mutualisms with other species, in which

both species typically benefit.
Natural selection mediated by
species interactions
• Selection mediated by species interactions is qualitatively
different from selection mediated by the abiotic environment.
• The factors in the environment causing selection are evolving

in response to adaptations in the focal organism – “adapting
to a moving target”.
• Another way to phrase this reciprocity, is that the relative

fitness of genotypes in one species is context-dependent and
the context itself can also evolve - G x G interactions.
Co-evolution
• As a result of this reciprocity, adaptive changes
in the traits of one species may trigger subsequent adaptive
changes in the traits of a second species, which in turn feedback to
cause further adaptive evolution of the first species.
• This process of reciprocal evolutionary change in interacting

species driven by natural selection, is called coevolution.
• Coevolution shapes interactions between pairs of species, small

groups of species, and large webs of species. It also affects how
species radiate into groups of descendent species.
• As the number of potentially coevolving species increases, so does

the number of ways in which coevolution can shape an interaction.
Co-evolution between plants and
antagonistic and mutualistic animals
• Plants adapt to animals in two main ways.
• First, they try to defend themselves from being eaten. This

defense can take many forms; resistance, avoidance, escape or
tolerance.
• These adaptations result in counter-adaptations in animals, to

circumvent the defense strategies, e.g. tolerating toxic
substances in plants.
Co-evolution between plants and
antagonistic and mutualistic animals ctd.
• Second, plants adapt by providing rewards for animals that carry out
beneficial services, such as nectar or pollen to pollinators, and fleshy
fruit pulp to dispersers.
• Plants, being sedentary, not only need to provide rewards to pollinators

and dispersers, but also need to make themselves detectable to
mutualistic partners, and evolve mechanisms to facilitate pollen and
seed transfer.
• Animals adapt to increase the efficiency in finding and using plant

rewards.
• This leads to a continuous sequence of adaptations and counter-

adaptations. This is equally true for mutualistic and antagonistic
interactions.
G x G interactions; e.g. hummingbirds and
hummingbird-pollinated plants
Co-evolution and diversity
• Much of the diversity on earth is a consequence of

coevolutionary diversification in species
interactions
Three important features
of co-evolution
1. Much of co-evolution is diffuse
2. Species interactions are not
stable over time
• Species interactions may change over time because the

identity of species change.
• Or, because the outcome of a given interaction changes.
• Over evolutionary time or over environmental gradients,

the outcome of a given interaction may vary and go from
antagonistic to mutualistic.
3. Interacting organisms have conflicting
interests, also in mutualisms
• One reason why the identity and outcome of interactions change
continuously over time, is that there is a fundamental conflict of
interest between interacting species.
• For antagonisms, like predation, herbivory and competition, this
conflict is obvious.
• However, also in mutualistic interactions there is

an underlying conflict of interest. Each partner
tries to gain as much as possible while contributing
as little as possible.
• Mutualisms can thus be

regarded as mutual
exploitation.
Structure:
interactions
selection
• Trade-offs
How to study natural selection
mediated by species interactions?
• Although, co-evolution is always a reciprocal

process, it is often convenient to study it from the
perspective of each of the partner.
Natural selection
• What are the essential components of evolution

through natural selection, as first suggested by
Darwin and Wallace?
1. Phenotypic trait variation
2. Fitness differences, as a result
of phenotypic variation
Fitness
Trait value
Heritability
Structure:
interactions
selection
• Trade-offs
Selection analyses
• So how can we study natural selection?
• Let us start with phenotypic selection and within-generation

changes, and then come back to the question about the sources
of phenotypic variability, and to the question of to what extent
effects are transferred to the next generation.
• Let us assume that we want to go out into the field and

investigate if there is natural selection.
• For example, investigating if there is selection on flower size in a

natural population of a herb.
• Then how can we identify and quantify selection?

Selection analyses
For example:
• Mark a random sample of plants
• Measure traits – flower size
• Measure fitness – seed number …
• Statistical analysis to explore the relationship – which?

Selection analyses – selection
differentials and selection gradients
• Analyses to explore relationships between traits and

fitness, i.e., to identify phenotypic selection, are well
described in Conner & Hartl.
• Read this chapter carefully, if you have not done so

already!
Selection differentials
Simple regressions of
standardized traits on
relative fitness
Relative fitness
Standardized trait value

Directional selection
Relative fitness

Stabilizing selection
Relative fitness

Disruptive selection
Relative fitness

No selection
Relative fitness

Selection differentials
Potential problems with inferring
natural selection and a causal
Relative fitness
relationship from this

relationship?

Targets of selection and Indirect
selection
Trait A
Fitness
Trait B
Selection differentials do not separate direct selection on a trait

from selection generated by effects of correlated traits on fitness
How can we deal with this problem?
Selection gradient analysis
• Multiple regressions of several standardized traits

on relative fitness
• Estimates direct selection, if all relevant traits are
included
• Additional problems?
Environmental co-variation
Environmental factors influence the

condition of individuals, which in turn
influence both trait values and fitness
How can we deal with this problem?
Trait A
Condition
Fitness
Structure:
interactions
selection
• Trade-offs
When does phenotypic selection
lead to evolution?
Relative fitness

Mean traait value of
offspring
Heritability
Mean trait value of parents

How can we investigate whether
observed phenotypic variation has a
genetic basis?
Mean traait value of
offspring
Mean trait value of parents

Formalizing heritability
• Vp = Vg + Ve
• Broad sense h2 = Vg / Vp
• Vg = Va + Vd + Vi
• Narrow sense h2 = Va / Vp
• Δz = h2 × S
• Vp = total phenotypic variation
• Vg = genetically based variation
• Ve = environmentally induced variation
• Va = additive variance; Vd = dominance variance; Vi = epistatic variance
• Δz = response to selection - trait change
• h2 = heritability
• S = selection differential - the average difference between the parent generation and
the selected parents
Measuring heritability
• Does it matter where and under what

conditions we measure heritability?
• Why?
Plasticity and Reaction norms
• Plasticity can be described as the ability of a given genotype

to be expressed as different phenotypes in different
environments
• Vp = Vg + Ve
• Patterns of plasticity can differ among individuals
• Vp = Vg + Ve + Vg×E
• Reaction norms are the pattern of phenotypic trait variation
for a given genotype along an environmental gradient, i.e.,
the trait value as a function of an environmental driver
Reaction norms describe how the expression
of traits depend on the condition of the
organism, or on the environment
e.g. flower size
Trait value –
Condition–
e.g. some measure of plant resource state
Reaction norms
e.g. development rate
Trait value –
Environmental driver –
e.g. some measure of temperature, integrated over time
Targets of selection
• Many, or even most, traits might be best be

described as reaction norms.
• For these traits, the reaction norm rather
than some specific trait value is the target of
natural selection.
Reaction norms differ among
individuals, and can evolve in
response to natural selection
A
Development rate
Temperature
Plastic trait responses are
changes along the reaction norm
Development rate
Temperature
Evolutionary responses are
changes in the elevation or slope
of the reaction norm
Development rate
Temperature
Mean reaction norms differ
among species
A
Development rate
Temperature
Differences in reaction norms
between interacting species and
environmental variation
A
Development rate
Temperature
Structure:
interactions
selection
• Trade-offs
Trait B (e.g. growth)
Trade-offs
Trait A (e.g. chemical defense)

Growth
Trade-offs
Chemical defense
Probability to escape predation/herbivory
Trade-offs
Chemical defense
Fitness component:
Trade-offs
Chemical defense
Fitness component:
Growth
Trade-offs – trait optima
Growth
Chemical defense
Optimum
Trade-offs
• Trade-offs play out differently in different
environments
• Let us assume that the example we just explored
was an environment with abundant herbivores
Chemical defense
Optimum
• How would the optimal level of defense change in

an environment with fewer herbivores?
Optimal defense when herbivores
Growth are rare
Chemical defense
are rare
Chemical defense
are rare
Optimal defense when

herbivores are abundant
Growth
Chemical defense
Optimum
Chemical defense
Optimum
Structure:
interactions
selection
• Trade-offs
Agents of selection
Trait Fitness
Environmental factor
– Selective agent
Agents of selection
Trait Fitness
Environmental factor
– Selective agent
How can we identify the
agents of selection?
Experiments –
e.g. pollinator-driven selection
Hand-pollinated
Controls
Fitness
Trait value (e.g. flower size)

Experiments –
herbivore-driven selection
Herbivores excluded
Controls
Fitness
Trait value (e.g. amount of chemical defense)

Experiments – herbivore-driven
selection
Herbivores excluded
Controls
Fitness
Trait value (e.g. amount of chemical defense)

Agents of selection
• The causes of natural selection are not correlations
between traits and fitness, but natural selection is
driven by the environmental factors that causes
differences in fitness between individuals with
differences in traits.
• This is an effect of the G × E interaction on fitness
• What are the implications of that the environment

varies over time and space for natural selection?
Structure:
interactions
selection
• Trade-offs
What do we mean by local adaptation?
What causes local adaptation?
Local adaptation is a mechanism whereby a

population of organisms evolves to be more well-
suited to its local environment than other members
of the same species that live elsewhere.
Local adaptation requires that different populations

of the same species experience different natural
selection and that gene flow is restricted.
Optimal defense in populations differing in
intensity of herbivory
Herbivores are relatively rare
Herbivores are abundant

Growth
Chemical defense
Optimum
Chemical defense
Optimum
Local adaptation in plant defense in two
populations differing in intensity of herbivory
are abundant
Fitness
Growth
Defense Intensity of herbivory
Fitness
Optimal defense when herbivores are rare A
B
Intensity of herbivory
Growth
Fitness
B
Defense Intensity of herbivory
Local population adaptation
• G x E effects on fitness mean that organisms can evolve traits that are
favorable in one environment regardless of how adaptive they are in
other environments.
• Selection and the optimal phenotype varies with the environment, and
the environment and selective agents varies spatially.
• Different local populations experience different selection and the optimal
phenotype differs among populations of a given species.
• Where such differences in selection are present and persistent, and
where gene flow is comparatively low, we expect populations to
differentiate genetically and become adapted to their local environment.
• This is what we call “Local adaptation”
• Several factors in the environment can cause local adaptation, not the
least differences in species interactions.
Local population adaptation
Sobral et al. 2015

How can we demonstrate
local adaptation?
Structure:
interactions
selection
• Trade-offs
Combining co-evolution with environmental
variation and local adaptation –
Geographic Mosaic Theory of Co-evolution
• Species do not interact as species, but as populations

• As a result, the evolutionary outcome of an interaction
between two species can vary over space, because:
 Populations differ genetically, e.g. due to local adaptation
 Abiotic environmental conditions differ
 Community context differ, e.g. in terms of the presence of
other herbivores, alternative pollinators or host plants
Geographic Mosaic Theory of
Co-evolution
Interacting with
genotype A of a different
species, or interacting in
environment A
Fitness
Interacting with
genotype B of a different
species, or interacting in
environment B
Trait level
Co-evolution
… argues that coevolution proceeds by natural selection acting
on three sources of variation that affect interactions among
species.
These three sources of variation can be formally partitioned as
genotype by genotype by environment interactions (G x G x E).
… proposes that coevolution exhibits three inherently
geographic characteristics, beyond local coevolutionary
selection:
1. Coevolutionary hotspots and coldspots
2. Geographic selection mosaics
3. Trait remixing
Coevolutionary hotspots
• Coevolutionary hotspots are communities in which the
pairwise interaction between two species exhibits
coevolutionary selection (i.e., there is a G x G
interaction on the fitness of both species).
• These hotspots are expected to be embedded in a

geographic matrix of coldspots. In coldspots, selection
is not reciprocal, including communities in which only
one of the species occurs (i.e., structural coldspots).
Geographic selection mosaics
• Geographic selection mosaics occur when natural
selection mediated by interactions differs among
environments (e.g., high vs. low temperatures, species-
rich vs. species poor).
• This variation occurs because the effects of one species
on the fitness of another species depends on the
environment in which the interaction occurs
interactions.
• That is, the fitness function of coevolutionary selection
differs among environments - G x G x E interactions.
Trait remixing
• The genetic structure of coevolving species continually
changes through new mutations, genomic alterations, gene
flow among populations, differential random genetic drift
among populations, and extinction of local populations.
• New genetic material on which natural selection can act can
result from simple genetic mutations, chromosomal
rearrangements, hybridization between populations, or
whole genome duplication.
• These processes contribute to the shifting geographic
mosaic of coevolution by continually altering the spatial
distributions of potentially coevolving genes and traits.
Co-evolution
Alternative hypotheses on the relative importance of the three elements of the

geographic mosaic theory of coevolution (GMTC) and non-reciprocal selection.
Co-evolution
Greya moths act as both

pollinators and seed parasites
of Lithophragma plants
Summary:
interactions are fundamentally important
• Co-evolutionary processes can be studied by considering each
partner separately
• In such analyses, it is crucial to identify the targets of selection
and the agents of selection
• Many traits influencing the outcome of species interactions are
plastic and can be best be described as reaction norms
• Trade-offs are fundamental to understand (co-)evolutionary
processes
• Local population adaptation and spatial structured species
interactions are fundamental and constitute the basis for the
“Geographic mosaic theory of coevolution”

Natural Selection and The Co-Evolution of Interactions

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Natural selection and the

• Both take place at the level of individuals

• Ecological effects on mean absolute fitness of individuals leads to

• Interactions can have an evolutionary but not an ecological effect,

• Natural selection mediated by species interactions and co-evolution is

• Most species compete with similar species

• Most species interact with other species as consumers (e.g. predators,

• Many species are engaged in mutualisms with other species, in which

• The factors in the environment causing selection are evolving

• Another way to phrase this reciprocity, is that the relative

• This process of reciprocal evolutionary change in interacting

• Coevolution shapes interactions between pairs of species, small

• As the number of potentially coevolving species increases, so does

• Plants adapt to animals in two main ways.

• First, they try to defend themselves from being eaten. This

• These adaptations result in counter-adaptations in animals, to

• Plants, being sedentary, not only need to provide rewards to pollinators

• Animals adapt to increase the efficiency in finding and using plant

• This leads to a continuous sequence of adaptations and counter-

• Much of the diversity on earth is a consequence of

• Species interactions may change over time because the

• Or, because the outcome of a given interaction changes.

• Over evolutionary time or over environmental gradients,

• However, also in mutualistic interactions there is

• Mutualisms can thus be

• Although, co-evolution is always a reciprocal

• What are the essential components of evolution

• Let us start with phenotypic selection and within-generation

• Let us assume that we want to go out into the field and

• For example, investigating if there is selection on flower size in a

• Then how can we identify and quantify selection?

• Mark a random sample of plants

• Measure traits – flower size

• Measure fitness – seed number …

• Statistical analysis to explore the relationship – which?

• Analyses to explore relationships between traits and

• Read this chapter carefully, if you have not done so

Standardized trait value

Standardized trait value

Standardized trait value

Standardized trait value

Standardized trait value

relationship from this

Standardized trait value

Selection differentials do not separate direct selection on a trait

• Multiple regressions of several standardized traits

Environmental factors influence the

Standardized trait value

Mean trait value of parents

Mean trait value of parents

• Does it matter where and under what

• Plasticity can be described as the ability of a given genotype

• Many, or even most, traits might be best be

Trait A (e.g. chemical defense)

• How would the optimal level of defense change in

Optimal defense when

Trait value (e.g. flower size)

Trait value (e.g. amount of chemical defense)

Trait value (e.g. amount of chemical defense)

• This is an effect of the G × E interaction on fitness

• What are the implications of that the environment

Local adaptation is a mechanism whereby a

Local adaptation requires that different populations