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Testing adaptive hypotheses Testing adaptive hypotheses

What is (an) adaptation? What is (an) adaptation?

Generally speaking, adaptations are traits or

characters that appear to be too well-fitted to
A trait, or integrated set of traits, that increases their environment to have arisen by chance.
the fitness of an organism. That is, they must be the result of selection.
The process of improving the fit of phenotype to Adaptations may involve morphological,
environment through natural selection physiological or behavioural traits. They arise
through the accumulation of a series of small
improvements over time.

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Testing adaptive hypotheses Testing adaptive hypotheses
What is (an) adaptation? Examples of adaptations

"If it could be demonstrated that

any complex organ existed The eye
which could not possibly have
Bird wings
been formed by numerous
successive slight modifications, The human brain
my theory would absolutely Homeothermic temperature regulation
break down." — Darwin Human language
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Testing adaptive hypotheses Testing adaptive hypotheses
Identifying adaptations Identifying adaptations – an incomplete example
Why do polar bears have white coats?
• Adaptive hypothesis: white coat is an adaptation for
In order to identify a trait as an adaptation, we camouflage
must first hypothesize its use or function, and • Test:
observe hunting behaviour and assay use of
then test that hypothesis. camouflage
As we saw in the Gould & Lewontin article, it is • Result: camouflage not usually important in hunting
important to test the hypothesis of adaptation • New adaptive hypothesis: white coat is an adaptation for
trapping solar heat
against a variety of null and alternative
hypotheses. • Test: hairs are actually clear and translucent, and trap 16%
of incident light energy – better than most hair types.
Results don’t support our first adaptive hypothesis.

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Testing adaptive hypotheses Testing adaptive hypotheses
Identifying adaptations – an incomplete example Identifying adaptations – an incomplete example
Why do polar bears have white coats? Why do polar bears have white coats?
• Adaptive hypothesis: white coat is an adaptation for • Adaptive hypothesis: white coat is an adaptation for
camouflage camouflage
• Test:
observe hunting behaviour and assay use of • Test:
observe hunting behaviour and assay use of
camouflage camouflage
• Result: camouflage not usually important in hunting • Result: camouflage not usually important in hunting
• New adaptive hypothesis: white coat is an adaptation for • New adaptive hypothesis: white coat is an adaptation for
trapping solar heat trapping solar heat
• Test: hairs are actually clear and translucent, and trap 16% • Test: hairs are actually clear and translucent, and trap 16%
of incident light energy – better than most hair types. of incident light energy – better than most hair types.
Results are consistent with our new adaptive hypothesis. What’s missing?
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Testing adaptive hypotheses Testing adaptive hypotheses
Identifying adaptations Approaches to testing adaptive hypotheses
Ex per im ent al st udies
The polar bear example shows that ecological and There are various experimental and theoretical ways to
physiological patterns are consistent with one particular Ex . test
Wing marks &ofwing
hypotheses waving
adaptation. inhas
Each Tephritid flies.
its benefits and
adaptive hypothesis – but it doesn’t show that fur color drawbacks:
evolved via a process of adaptation. • Observational studies (e.g., the polar bear study)
We’ve compared alternative hypotheses of adaptation – Tephritid flies have dark bands on their wings and
• Experiments
wave their wings when disturbed in a manner that is
but we haven’t tested the biological null hypothesis:
that no adaptation has occurred. • Theoretical
reminiscent of their models
major predator's territorial
The primary null hypothesis is that traits have evolved display• --
Comparative method
e.g., jumping spiders' leg waving.
due to drift (according to the neutral model). • Molecular evidence
Do flies mimic their predators?
If so, is this mimicry to deter any predator, or is it
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specifically to deter jumping spiders? 12
Testing adaptive hypotheses Testing adaptive hypotheses
Experimental approaches: example Experimental approaches: example

Tephritid flies have dark bands on their wings. When

disturbed, they wave their wings in a manner
reminiscent of the territorial behavior of their predator
(leg waving in jumping spiders).
Do the flies mimic their predators?
Does this mimicry deter predators?
Does it deter jumping spiders in particular?
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Testing Adaptive Hypotheses
Text
Conclusion:
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Testing adaptive hypotheses
Theoretical approaches
Two classes of models predict how a trait should evolve
under a specific set of environmental circumstances
(usually ignoring genetics altogether).
• Optimality models assume that a trait will evolve
to impart the highest possible fitness.
• Evolutionarily Stable Strategy (ESS) models assume
that the fitness of a phenotype depends on what
other phenotypes are present. This doesn’t always
lead to maximal fitness.
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Testing adaptive hypotheses
Experimental approaches: example
The evidence suggests that the morphology and
behavior of the flies increases their fitness in the face of
their primary predator.
Unlike the observational approach, this gives direct
evidence of a fitness advantage.
However, experiments like this still don’t directly test
the hypothesis that the traits evolved due to the
process of adaptation.
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Testing adaptive hypotheses
Theoretical approaches – example
We might do an ESS model to explore the possibility
that lekking behavior in birds is an adaptation to
predation risk (because the risk of predation gets
spread out among large numbers of males).
The model would predict the number of males we
should see on a lek as a function of the risk of
predation and the likelihood of mating.
We could then measure whether the number of males
we observe in nature is consistent with the predictions
of the model, given our observations of predation and
mating probabilities.
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Testing adaptive hypotheses Testing adaptive hypotheses
Theoretical approaches – example The comparative method

These models offer quantitative predictions for

observational or experimental studies.
However, they seldom compare predictions to
alternative hypotheses (either adaptive or non-adaptive)
Also, the quantitative predictions might not take into
account constraints on adaptation – any failure of
observations to match predictions might be due to such
constraints.
If we have an adaptive hypothesis for a trait, then we
might expect to see a correlation between the
explanatory variable (X) and the trait itself (Y).
However, another possible explanation for such a
correlation is the process of evolution itself: organisms
whose common ancestors had both X and Y are also
Thelikely
Comparative Method
to have X and Y.
Ex . Testis size in bat species.
Male bats vary in the size of testes. One hypothesis
for this variation is that large testes produce more
sperm, an advantage in sperm competition.

19 Sperm competition might be more intense in larger 20

Testing adaptive hypotheses Testing adaptive hypotheses
social groups, where more males compete for
reproductive access to females.
The comparative method – Example The comparative method – Example
Prediction:
Hypot hesis:Species
Testis with
size islarger
largersocial groups
in species withshould
Testis size in bats have males
larger with
social larger testes (because more males are
groups.
Male bats vary from species to species in the size of competing for reproductive access to females).
their testes.
Adaptive hypothesis: Larger testes produce more
sperm, which provides an advantage if sperm from
multiple males competes for fertilization in a female.
Prediction: Species with larger social groups should
have males with larger testes (because more males are
competing for reproductive access to females).
 1. Need phylogeny
2. Calculate contrasts between sister taxa
Closely related3.species
Evaluate relationship with phylogenetically-
21 might have similarcorrected
group 22
Testing adaptive hypotheses Testing adaptive hypotheses
values.
size
But not all data points are and testis size NOT
independent. because of sperm
The comparative method – Example The comparative method – Example
competition, but because
they share an ancestor
Null hypothesis: Closely related species might have who had large testes and
similar group size and testis size simply because they
Closely related species lived in large groups.
share a common ancestor.might have similar group
size and testis size NOT
But not all data pointsbecause
are of sperm
independent. competition, but because How do we control for
they share an ancestor the effects of shared
who had large testes and history?
lived in large groups.
Closely related species
might have similar group
size and testis size NOT
because of sperm How do we control for
the effects of shared
competition, but because
they share an ancestor history?
who had large testes and
lived in large groups.
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Testing adaptive hypotheses Testing adaptive hypotheses
The comparative method – Example The comparative method
How do we control for
Controlling for common ancestry,
the effects males in species
of shared with Evidence that phylogenetically independent contrasts of
larger groups sizes still have larger testes.
history? a trait (e.g., testis size) are correlated with a
hypothesized explanatory variable (e.g., group size)
suggests that:
• The trait has evolved in the absence of (or despite)
phylogenetic constraint
• Not all evolution of the trait has been neutral
(because there is evidence of directionality)
The comparative method explicitly tests adaptive
hypotheses against a null hypothesis.

Yes, males in species with larger group sizes also

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Testing adaptive hypotheses
The comparative method – caveats

“Correlation is not causation.”

Still not a direct rejection of the neutral model.