15/02/2017

Behavioural Ecology

Chapter 1
An evolutionary approach to
animal behaviour

Robby Stoks

Behaviour ?

“All observable processes by which an animal responds

to perceived changes in the internal state of its body or
in the external world.” (Skinner)

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Pioneers in the study of animal behaviour

• Nobel prize in Medicine in 1973

Niko Tinbergen Konrad Lorenz Karl von Frisch

(the Netherlands) (Austria) (Germany)

Scientific Method
• Tinbergen used the scientific method (question,
hypothesis, prediction, test, and conclusion) to
understand beewolf homing behaviour.

• Behaviour: When leaving nest to hunt bees, female

beewolf covers nest entrance with sand

• Question: How do female beewolves find their way

home?

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Scientific Method
• Observation: Beewolves circle next before leaving.

• Hypothesis: Beewolves use landmarks to remember their

nest hole.

• Prediction: When landmarks are removed

beewolf should have difficulties relocating her nest.

• Test 1: Removal of objects around nest => females

struggled to relocate nest

Scientific Method
• Test 2: Setup landmarks around nest to use, then move
landmarks => female searched where landmarks suggested
nest should be.

• Conclusion: female bee wolves use landmarks to relocate

their nests.

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Tinbergen’s Four Questions

• (1) Development:
How does the behaviour develop? Is it learned? Is it present at
birth? Which genes turned on/off?

• (2) Mechanism:
How is the behaviour achieved? How does an animal uses its
sensory and motory abilities to activate and modify its behaviour
patterns?

• (3) Function:
How does the behaviour promote survival and reproductive
success?

• (4) Evolution:
How did the behaviour evolve? How does an animal’s behaviour
compare with that of other related species? Was it present in the
ancestor? What are the intermediate steps in the evolution of the
behaviour?

Tinbergen’s Four Questions

• (1) Development
Proximate
Ethology

• (2) Mechanism
Behavioural

• (3) Function
Ultimate
ecology

• (4) Evolution

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Tinbergen’s Four Questions

Darwinian theory and ultimate

hypotheses

Evolutionary change in a trait is inevitable if following three conditions are met:

1. Variation, such that members of a species differ in the trait.

2. Heredity (i.e. genetic variation), with parents able to pass on their value of
the trait to their offspring.

3. Differences in reproductive success, such that some individuals have more

surviving offspring than others in their population thanks to their trait value.

Process that causes evolutionary change is natural selection.

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Darwinian theory and ultimate hypotheses

1. Genes code for proteins that will eventually shape traits within a species.

2. Genes may exist in two or more forms, or alleles, within the species’ gene pool.
Different alleles may code for slightly different proteins.

3. Competition between alleles at a given locus on the chromosomes.

4. Allele that makes more copies (higher reproductive success of its carrier) will
gradually displace its competing alleles.
Natural selection = differential copy succes of alternative alleles.
Evolution = change in allele frequencies in a population through time.

Unit of selection = gene (individual = gene copying machine) (Dawkins)

Example of behaviour that one can only explain by reasoning at the gene level?

Group selection

• Unit of selection = group

• V.C. Wynne Edwards (1962): Individuals behave in the interest of their

group, often at a cost to themselves

• Group selection chooses between groups on the basis of how well their
individuals serve the group’s interests, i.e. increase the ability of the group
to survive.

• Examples:
- Males compete for females to ensure only the strongest can reproduce
which is best for the group.
- Honey bees sacrifice their own reproduction for the good of the colony.
- Infanticide occurs to keep densities low and not to overexploit resources.

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Problems with group selection

Problems with group selection

Thought experiment:
• Imagine a limited amount of food for a population. Group selection
might produce animals that voluntarily restrict food intake “for the
good of the group”.

• A mutation occurs for eating more than ones share (“mutant

cheater”)
– More food intake = more reproduction

• Mutant increases in allele frequency and approaches fixation.

• Natural selection favors selfish cheaters.

– Therefore, Cheaters Always Win.
– Altruism and group selection generally do not win.
– Notable exceptions are things like kin selection
(to be discussed later).

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Group selection
• Current view:
- Group selection may work
- Most likely when groups are isolated from each other
and differ genetically in ways that affect their chances to
survive locally.
- Group selection is, however, likely much less important
than natural selection at the individual/gene level.

How to explain infanticide?

• Problem: infanticide does not seem to increase an
individual’s reproductive success. Moreover, risk of
being wounded.
• Adaptive value?

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How to explain infanticide?

Langurs live in groups of one male, several females
and their young.

Males fight to control groups of females.

When new male takes over group, infants

often killed by the new male.

How to explain infanticide?

• Social Pathology (SP) hypothesis
Males in high density populations become hyper-aggressive and so attack
babies.

• Quicker Reproduction (QR) hypothesis

When her nursing infant is killed, mother might resume her reproductive
cycle sooner than otherwise, and so could be impregnated sooner by the
infant killer, who leave more descendants as a result.

• Cannibalism (C) hypothesis

Males kill infants after taking over a band of females to eat the dead
youngsters, thereby replenishing reserves depleted in the takeover struggle.

• Population Regulation (PR) hypothesis

Self-sacrificing males take the risk of killing infants to avoid overcrowding
which would otherwise result in depletion of resources and population
extinction.

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How to explain infanticide?

• Infanticide does not occur more at high densities.
=> against SP and PR hypothesis
• Infanticide occurs soon after group has been taken over
=> supports QR and C hypothesis
• Males do not consume infants killed
=> against C hypothesis
• Males do not kill their own babies
=> supports QR hypothesis
• Females promptly resume sexual recycling and become receptive to
the male that killed their youngsters
=> supports QR hypothesis
• Infanticide also in other species with similar social system
=> supports QR hypothesis

How to explain infanticide?

Jacanas: Polyandrous (one female mates
with several males).

Female taking over a territory kills any chicks of

previous female. Frees up male to tend new female’s
eggs.

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