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psyc 357 notes

evolutionary background
evolution
all living things have descended from common ancestors (evolution as history) by a natural
historical process of change and diversification (evolution as process)

darwin's three concepts

1. what evolution produces: adaptation
2. evolution as history: descent with modification (phylogeny)
3. the mechanism of evolution: natural selection

(1) adaptation
characteristics of a species are adapted to its ecological niche
• (the way the species “makes its living”, i.e., where it lives, what it eats, what eats it, etc)
mimicry as example:
blending to your background or something other than what you are
• behavior is relevant – the animal has to put itself on the right background!
• mimicry occurs in all sensory channels
defensive mimicry: for the purpose of avoiding or escaping from your predators
aggressive mimicry: for the purpose of capturing your prey

(2) descent with modification (phylogeny)

any 2 species can be traced back ultimately to a common ancestor
the differences between them are the result of the history of the adaptations of their respective
lineages since they diverged from their common ancestor!

communication signals as example of historical approach: threat signals

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open mouth thread (uakari, yellow baboon, bushbaby) & tense-mouth thread
phylogeny of communication signals/displays
kessel (1955): comparative study of nuptial displays in many species of empid flies (including h.
sartor)

(in the second species, male presents female a nuptial gift - a smaller fly)
phylogeny of nuptial gifts in empid flies:

phylogenetic hypothesis of the evolution of the balloon display:

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(3) natural selection

distinguish between
evolution as history
• the idea that different species have descended from a common ancestor, and
evolution as process
• how different species come about
• how animals become adapted to their environment
darwin: evolution comes about through natural selection (a process analogous to artificial
selection)
natural selection
1. the struggle for existence
• reproductive rates are very high
• but most populations are stable
• why? – natural checks and balances: predators, parasites, limited resources
2. the mechanism: natural selection
• individuals vary within a species
• variation is in part heritable
• variation is related to adaptation
thus, if individuals vary re some trait related to reproductive success, the better adapted
individuals will leave more offspring and next generation will be different (better adapted).
natural selection occurring over a long time will lead to dramatic changes in a species - to
evolution
example: giraffe ecological niche = a high browser

okapi – giraffe’s closest relative

ancestral species (probably) was like the okapi (with a regular-sized neck)
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over time: a1 alleles will replace a2 alleles in the population and giraffe necks will get longer
(natural selection)
natural selection = differential reproductive success (RS) of individuals within a population due
to genetic differences among them
will illustrate the importance of each in following examples involving red deer
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reproductive success (RS), not survival, is the “bottom line” for natural selection
questions:
• has natural selection occurred if A and C are identical twins?
• (not twins) has natural selection occurred if A was killed by A lightning bolt?
• has natural selection occurred if the key differences between A and C are based on
learning? (assume they are unrelated)
natural selection = differential reproductive success (RS) of individuals within a population due
to genetic differences among them

some important implications and qualifications

1. if no genetic basis, then no natural selection
2. reproductive success (not adaptation, not survival) is bottom line for natural selection
3. selection acts on individuals (not species, not groups, not populations)

selection generally acts on individuals (not species, not groups, not populations):
modern study of animal behavior coincides with the realization that selection acts at the level of
the individual (or the gene actually) and rarely at the level of larger groups (group selection).
two problems with group selection thinking:
• logical problem (the problem of altruism)
• generally fails to predict correctly

the logical problem with group selection

lemmings are small rodents that live in the arctic tundra — at high population densities, large
number leave their homes and travel long distances, during which time many die, some by
drowning, as they attempt to cross rivers and lakes
• one popular explanation for their behavior is that the travelers are committing suicide to
relieve overpopulation
by heading off to die, the suicidal lemmings leave shelter and food for those that stay behind;
and these surviving individuals will perpetuate the species, saving it from extinction
what’s the problem with this hypothesis?
the logical problem with group selection is that if the difference between suicidal behavior and
nonsuicidal behavior is based (even only partially) on a genetic difference, then the non-suicidal
genotype (the lemming with the life preserver in this cartoon) will become more common in
future generations (will be naturally selected)!
so how can the ‘altruistic’ or suicidal genotype evolve?

 recognizing that selection acts at the level of individuals helps us understand how
behaviors that do not appear to be ‘adaptive’ might still be favored by natural selection
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evolution of infanticide in langurs

in this question you need to contrast three stages of evolution:
(1) before infanticide appeared,
(2) during the directional phase of selection (both infanticidal and non-infanticidal males in
population), and
(3) the present – all males assumed to be infanticidal
question: after infanticidal males first appeared in langurs, only one of the following effects on
fitness would have actually occurred — which one is that? (important note!: throughout
“offspring” means “offspring surviving through weaning”)
a. males would have more offspring than would females
b. the total number of offspring (say, per year) of the langur population as a whole would
increase
c. infanticidal males would leave more offspring than would non–infanticidal males
d. the average infanticidal male (now) would leave more offspring than the average male
before infanticide appeared in the species

feeding chases in adelie and emperor penguins

kids are parked in the ‘creche’ while both parents go to sea to feed — parents return to feed kid;
kid then has to chase parent to get fed!
question: why the feeding chase?
“it enables each returning adult to identify and feed the chicks most in need of food since the
hungriest will be most persistent — this feeding strategy has evolved through natural selection
to ensure that despite the leopard seals most penguin chicks will get enough to eat”

group selection & “altruism”

selection acts on individuals (not species, not groups, not populations)
but many behaviors do appear to benefit the group at a cost to the individual: these are called
altruistic behaviors
altruism: an individual acts to further the reproductive success of another individual(s) at some
(reproductive) cost to itself

 example: belding’s ground squirrel alarm calls

group selection and altruism: the prediction problem

belding’s ground squirrels
• alarm calling is valuable to unaware ground squirrel that hears it but potentially risky
(exposes caller to danger) = “altruism”
tioga pass, sierra nevada mountains, california
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research by paul sherman & an army of undergrads

they tested whether alarm call is risky — are callers more likely to be taken by the predator than
are non-callers?
belding’s ground squirrel has two kinds of alarm calls. terrestrial alarm call (trill) given to
coyotes

aerial alarm call (whistle) given to hawks:

need to know about these animals:

• parental care by females only
• males provide sperm, that’s it
individual and group selection arguments make different predictions as to whether these classes
of animals should alarm call:
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class group selection prediction individual selection prediction

males yes no? (depends on whether other
relatives or present)
females without pup yes? no/yes (depends on whether
other relatives or present)
females with pups no yes
pups no yes

need to know about social organization:

• parental care by females only
• males disperse from natal group, females stay
• males also move away during breeding season

battle at kruger
african (cape) buffalo

 male ~ 5 – 5.5 ft, 1300 – 1700 lbs

 female ~ 4.5 – 5 ft, 900 – 1200 lbs
both have heavy, ridged horns
large, mixed-sex & -age herds, with exclusive home ranges
basic unit within herd = clan of 12 or more related cows & their offspring
3 to 5 breeding (i.e., dominant) bulls consistently accompany each clan
bachelor herds: bulls past their prime, young males (3-8 years old)

— why do the buffalo take so long to attack the lions?

— why do they eventually attack? who do you think attacks the lions first? the strongest
buffalo in the herd? one of the dominant bulls? the weakest/most expendable buffalo?
mom? dad?
each buffalo is initially trying to save his/her skin, largely by trying to stay with the herd. lions
are cooperative hunters (i.e. hunt as a group), and while a single lion couldn’t take on a buffalo,
even perhaps a baby buffalo, if they manage to isolate one from the herd, they could take
him/her down — so the lions are trying to peel off a buffalo (preferably a young one without a
ridge)
after they manage to do that, the lions have to deal with the unexpected rival for their meal —
meanwhile the buffalo come back to see if they can save the calf. again for each individual
buffalo, the cost of attacking the lions first as an individual is potentially quite high, so you’d
prefer someone else to take the lead first
hence, they surround the lions but it takes them a while to get attacking
the first attacker likely is a relative, and mostly likely a female relative, the mother or a close
female relative: because there are multiple breeding males, there is some uncertainty with
respect to who the father might be, but the mother doesn’t have that uncertainty — so they have
the most benefit to gain from saving the calf, and should be more willing to take the higher risk
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(remember at the point of first attack, when the buffalo send a lion airborne, the entire pack is
still there, so that’s the most risky moment)
after the lions are routed, and they start retreating one-by-one, the chasing buffalos may indeed
be anyone at the herd: at point there isn’t much risk a single retreating lion won’t be too much of
a threat to an adult buffalo

types of questions about animal behavior

questions about behavior: different levels of analysis (different perspectives)

historically, in study of behavior, there has been confusion between explanations at different
levels (especially between proximate & ultimate)
but explanations at different levels are almost always complementary, not alternative
integrative explanations involve several levels

tinbergen’s “four questions” (perspectives)

1. cause or mechanism — proximate
a. environmental factors (stimuli)
b. inferred mechanisms (psychological)
c. genetic mechanisms
d. neuro-physiological mechanisms
2. development — proximate
3. function (how is it adaptive?) — ultimate
4. phylogeny (evolutionary history) — ultimate

• proximate (1 and 2)
• and ultimate (3 and 4)
explanations are considered to be different levels of explanation
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langur infanticide & levels of explanation

ultimate – functional: males who kill nursing infants after take-over have greater reproductive
success than males who do not
not a proximate explanation! in particular, not a hypothesis about the male’s motivation or
thought process
avoid confusing proximate & ultimate explanations!
not: “hmm, i better go kill an infant and increase my reproductive success…” we’re not saying
this! ultimate explanation is that male increases his RS by infanticide but who knows what he’s
thinking? — probably not this!
watch out for our use of the “language (metaphor) of decision”— it is a short-hand way of
expressing an ultimate (functional) cause as if it were a proximate cause.

“song” complexity as an example of proximate vs. ultimate approaches to a question

many animals have complex vocal signals they use as species identification signals to attract
mates and/or ward off competitors
background: many animals have complex vocal signals they use as species identification signals
to attract mates and/or ward off competitors
complex vocal signals are:

learned ‘hard-wired’
humans all other primates
cetaceans (whales & dolphins) all other mammals
bats (?)
songbirds, hummingbirds, parrots all other birds
all frogs
all fish
all insects (e.g. crickets)

bird song repertoires & mimicry

an example illustrating the differences among tinbergen’s “four questions” and the distinction
between proximate and ultimate explanations
example: in some songbirds, the male (in temperate-zone species, usually only the male sings; in
tropical species, female often sings as well) sings one species-specific song, in others, multiple
song types (aka a “song repertoire”), sometimes mimics other species, other sounds — why?

(a sparrow song)
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song mimicry
superb lyrebird
question: in some songbird species, birds sings 1 species-specific song, but in others, he sings
multiple song types (aka a “song repertoire”) and in some he even 'enlarges his repertoire' by
mimicking other species, other sounds
think of hypotheses of any of the four types (functional, phylogenetic, causal, or developmental)
to explain this phenomenon
1. cause or mechanism
a. environmental factors (stimuli)
b. inferred mechanisms (psychological)
c. genetic mechanisms
d. neuro-physiological mechanisms
2. development

3. function (how is it adaptive?)

4. phylogeny (evolutionary history)

two main general functions of song

1) female attraction
2) male competition

but how/why do song repertoires help? functional hypotheses

1) beau geste
2) female preference
3) ...many others...

krebs – beau geste hypothesis

territorial bird sings many songs to deceive prospecting birds into thinking there are (too) many
birds there and so to move on
what testable predictions follow from this hypothesis?
one test: removal – the keep-out experiment

 are repertoires more effective than a single song? how can we test?
repertoire of songs more effective as keep-out signal than is single song (both presented equal
number of times).
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female preference
hypothesis: females prefer hypothesis: females prefer
males with larger song males with larger song
repertoires repertoires
field observation: males lab: tests using copulation
with larger repertoire solicitation display –
sizes are chosen first by females solicit more to
females (sedge warblers) larger repertoire sizes
(various species)

ultimate – phylogeny (evolutionary history)

1. did song-learning evolve in common ancestor of all birds?
2. is learning a necessary precondition for repertoires to evolve?

1) did song-learning evolve in common

ancestor of all birds?

 no, it appears to evolved 3 times

independently
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2) is learning a necessary precondition for repertoires to evolve?

neuro-physiological mechanisms
(proximate – neural mechanisms)
proximate hypothesis: larger song-control
brain regions lead to larger song
repertoires’
finding: repertoire size and HVC size are
correlated (across species)

each data point represents a species

(actually a species contrast)
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furthermore: repertoire size and HVC size

are correlated within and between east and
west marsh wren populations

why is repertoire size larger in western marsh wrens than in eastern marsh wrens?

western marsh wrens have larger song repertoires (100-200 songs) than eastern marsh wrens
(~50)
hypothesis 1: differences in two populations are due to genetic differences
hypothesis 2: differences are due to western birds living in denser populations and thus hearing
more songs during the sensitive period ‘common garden’ experiment: bring chicks from both
populations into the lab, give them identical song tutoring regime
‘common garden’ experiment: bring chicks from both populations into the lab, give them
identical song tutoring regime: by hypothesis 1 resultant repertoire sizes should be different, by
hypothesis 2 the same

kroodsma’s anti-habituation hypothesis

differences in population density (W > E) lead to differences in repertoire size (W > E)
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song tutoring (‘common garden’) experiments

developmental/genetic mechanisms a ‘common garden’ experiment
why does a bird have more than one song type? what determines repertoire size?
proximate hypotheses:
• males with bigger song-control regions in the brain have bigger song repertoires
• repertoire size differences are (partly) genetically determined
• males who hear more different songs when they are young develop bigger song
repertoires
ultimate hypotheses:
• females prefer to mate with males that have more songs
• males with bigger repertoires do better in male-male competition
• vocal complexity (e.g., song repertoires) can only evolve after song learning evolves

an integrative hypothesis:
• does sexual selection for increased repertoire sizes lead to adult song learning (i.e. open-
ended learning)?
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satin bowerbird
(australia & new guinea)
a mimic:
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satin bowerbirds — good mimics have higher mating success (are more attractive to females?)

a scientific trade-off:
• ecological validity — resemblance of experimental situation to the real world
• separation and control of variables — ability to distinguish the effects of independent
variables X and Z on dependent variable Y (i.e., ‘internal’ validity)

ecological validity separation & control of variables

field obsevation best worst
comparative method best worst
field experiment better better
lab experiment worst best

optimality and comparative method

1. adaptations for survival
2. comparative method
3. cost-benefit and optimality approaches

adaptation, trade-offs and optimality

behaviors have
costs (C)
as well as benefits (B)

 a trait is adaptive if B > C

 a trait is optimal if B – C is maximal

say, long necks of giraffes have their

advantages (benefits: feeding) and their
disadvantages (costs: difficulty in drinking,
copulation)
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questions about adaptation:

1. is trait adaptive?
2. in what way is it adaptive?
3. is it optimal?
examples:
1. costs and benefits of calling in male field crickets
2. sperm competition in primates
3. parent-offspring recognition in swallows
4. optimal foraging
(2 and 3 are good examples of comparative method)

costs and benefits of cricket calling

a case where we can identify major costs and benefits
• benefit: attracting a mate
• cost 1: attracting parasitic flies
• cost 2: attracting satellite males

evolution of silent male crickets on hawaiian island of kauai

normal male wings have a toothy vein that is scraped to make sound — in mutant males, that
vein is smaller and repositioned
(females don't have this toothed vein at all)
the mutation that leads to the mutant male is a single mutation on the X chromosome (in the
crickets, the females have two X chromosomes, males have a single X chromosome and no Y
chromosome)
a single mutation in a sex-linked chromosome allows the mutation to spread rapidly in the face
of strong selection
the silent phenotype emerged on a different island as well (oahu)
evidently, the silent phenotype in oahu is caused by a different mutation (again on the X
chromosome)
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sperm competition
comparative approach
when multiple matings occur, increased sperm count provides an increased chance of
fertilization — a greater number of sperm are produced by larger testes
prediction: testes size should be larger in species with sperm competition (multiple male
matings) — test in primate species
how to test? compare primate species
1. monogamous (unusual) — gibbons, siamangs, owl monkeys, humans
2. single male groups — langurs, gorillas
3. multi-male groups — chimpanzees, most baboons
prediction: testes should be larger in species with multi-male groups than in either monogamous
or single-male group species
analysis of function based on comparison of convergent species — two or more species of
distant common ancestry that have similar traits because of the similarity of their ecological
niches or selection pressures

 convergent mating systems

 divergent mating systems

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allometry: how characteristics of organisms change with size

• full circle on the top right: chimp

• plus sign (+): humans
• triangle on the far right: gorilla
we can measure relative testes size = observed size / predicted size
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question: why the different pattern for dimorphism and canine size vs. testes size? — probably
related to dominance (intrasexual selection to be the ‘single-male’ with harem)

a second example of comparative method: swallows

similarities:
• aerial insectivores
• monogamous
• shared parental care
• monomorphic (mostly)
differences:
• nesting
• coloniality
• mate-guarding
• parent-offspring recognition

• bank swallows (colonial)

• cliff swallows (colonial)
• barn swallows (solitary)
• tree swallows (more or less solitary: when nest boxes placed in groups they have to be
placed at least 10 feet apart)

comparative method
analysis of function based on:
comparison of convergent species — two or more species of distant common ancestry that have
similar traits because of the similarity of their ecological niches or selection pressures
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OR divergent species — two or more species of recent common ancestry have dissimilar traits
because of the dissimilarity of their ecological niches or selection pressures

parent-offspring recognition in bank swallows

 bank & cliff convergent with respect to coloniality

 tree & barn convergent with respect to solitary living
adaptation

 bank & cliff convergent with respect to parent-offspring recognition

 tree & barn convergent with respect to lack of parent-offspring recognition

both parents feed kids at the nest for ~ 21 days, then for a week or so outside the nest (in
crèche)
• before kids leave nest for good, there is a transitional period during which they fly in and
out, sometimes flying back into wrong burrow
• on returning to the nest (burrow), parents will ‘evict’ unrelated young they find in their
burrow

playback test of parents’ recognition of their chicks’ calls

comparable tests with tree, cliff and barn swallows.

 bank swallows and cliff swallows show recognition in these tests, barn swallows and
tree swallows do not
parent-offspring recognition (discriminating parental care): cross-fostering experiments
trade 2 chicks from nest a with 2 chicks
from same-aged nest b, and leave 2 chicks in
each nest
(actually, remove and mark them in similar
fashion, then return them to home nest)
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answer:
the real phylogeny, because it shows that the species of distant common ancestry have similar
traits (i.e., they are convergent with respect to these traits) because of the similarity of their
ecological niches (colonial or solitary living)
the real phylogeny, then, provides a functional explanation: if it was the other way around, then
we could not infer whether their similar trait is due to their phylogeny or due to an adaptation to
colonial living

(optimal) foraging
foraging: searching for and exploiting food resources

types of foraging strategies

generalists vs. specialists

foraging strategies
• search
• ambush or sit-and-wait
• social foraging
• information sharing/extraction
• food storing (caching) — clark’s nutcracker etc.
• tool use — chimp etc.

optimal foraging problems

what to eat, or, what to forage for?
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elements of optimality approach

1. a set of decisions, choices or strategies
2. the currency, or the criterion used to compare the value of the alternative decisions
3. the constraints or limits (internal or external) on the animal
4. the prediction based on a theory or model
5. the test

an optimal foraging problem: the fox went out on a chilly night…

and he had to decide: should he forage

 in the town for the farmer’s grey goose?

 or in the fields for mice?
 or in the thickets for partridge?

given the facts below, and assuming the animal is trying to maximize its net rate of gain
(measured in grams), decide which would be the better choice:

prey type encounter rate prob. capture body wt (meat)

mouse 2 per hr ½ 50 g
partridge 1 per 10 hr 1/10 500 g

encounter rate multiplied by the probability of capture multiplied by the meat (grams)
= x grams/hour

mouse vs. partridge example

simplified in many respects
we assume:

 the only cost is time (but often there are energetic costs as well)
 the animal can hunt for only one or the other prey (but often it can hunt for two or more
prey types)
 the animal is selected to maximize the
net rate of energy gain = net intake / time required

the prey choice (diet selection) problem

what if the fox can hunt for partridge and mice at the same time? should he stop and eat
whichever one he encounters? or should he eat just the more profitable one? two kinds of prey,
1 and 2:
• energy value (calories) E1 and E2
• handling time h1 and h2 (constraint)
• encounter rate λ1 and λ2 (or search time S1 and S2)
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if E1 / h1 > E2 / h2 , prey type 1 is the more “profitable” prey

currency = net rate of energy intake
what should the fox do?

predictions of the prey choice model

(profitability)
animal should specialize (eat only more profitable prey, type 1) or generalize (eat type 1 and
type 2), i.e., exclusion of less profitable prey should be all-or-nothing
decision to generalize or specialize depends on S1, not S2:
you should generalize if

if > 2 prey types, types should be added to the diet in order of their profitability (i.e., according
to equation above, profitability)
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number of assumptions including:

• predator has complete information (does not need or use information it acquires while
foraging)
general point: handling less profitable item (prey 2) takes time that you could use searching for
(prey 1) — that’s why h2 is in equation but S2 is not!

a lab test of the prey choice model

the patch residence model

how long to stay in a patch, when to move on?
best examples: central place foragers (honeybees, chipmunks, monkeys, birds)

stay until the expected net energy gain in

the current patch drops to the same as the
gain from traveling to and foraging in a new
patch (marginal value theorem)
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how long to stay in a patch?

gain follows law of diminishing returns because:
1. the forager could deplete the patch
2. or could lose ability to carry food home

marginal value theorem graphical analysis:

law of diminishing returns

the tangent line to the curve gives you the

optimum

assumptions of patch model (MVT)

• forager has complete information (ex: knows travel time between patches)
• gain curve is negatively accelerated (i.e., diminishing returns)
• searching for and foraging within a patch have equal energy costs
• searching for and foraging within a patch are mutually exclusive activities

predictions of patch model (MVT)

• if travel time and the gain curve are known, then Topt can be predicted
• animal should stay longer in patch when travel time to patch is longer
• animal’s stay time is unaffected by patch quality (when patches uniform)

 animal should stay longer in patch when travel time to patch is longer
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animal’s stay time is unaffected by the patch quality (when patches uniform) — they gain less
amount of food in the lower quality patch compared to the higher quality patch, but the Topt is the
same

lab test of patch model

 patches = cups containing mealworms in sawdust

 patch quality = density of mealworms in sawdust
 travel time = tightness of lids on cups (more like handling time here, but effect is the
same)
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other ways of studying optimal foraging

foraging for whelks by NW crows
questions & answers:
what size whelks to take?

 only dropped large whelks (3.8-4.4 cm)

what height to fly to?

 flew to ~15 feet (5 m)

what to do if whelk doesn’t break?

 kept trying!

next question: are these behaviors optimal? how do you test?

test: built 15-m tower with adjustable platform, dropped different-sized whelks from different
heights
results:
1. on average, large whelks require fewer drops (also have more meat!)
2. probability large whelk will break increases as height increases, but changes very little
after 5 m

3. probability large whelk will break = 25-30% for any drop, independent of how many
times dropped
4. energy required to open average large whelk = 0.5 kcal, energy in = 2.0 kcal, net gain =
+1.5. medium whelks net gain = -0.3; small whelks worse

kleptoparasitism
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different goals and constraints

there are numerous factors that might cause animal to make a decision other than the simplest
optimal one of maximizing its rate of energy gain:
different goals:
• avoid predation
• maximize energetic efficiency (E gained / E spent)
• in variable environments, minimize the chance of starvation or maximize the chance of
survival
constraints:
• nutrient needs
• physiological constraints
• search image

balancing nutritional requirements

moose has to balance requirements for energy and sodium

 aquatic plants: rich in sodium, poor in energy

 terrestrial plants: rich in energy, ~ no sodium

physiological constraints

weddell seals:
90% of dives are within aerobic dive limit
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as the dive goes on, lactic acid builds up in

the muscles
(but not in the blood vessels, due to
vasoconstriction!)

by spending less time under water per dive, the weddell seal are able to spend more time under
water!

 if they dive under the ADL (about 20 minutes), they only need to replace the O 2 stores
 if they dive over the ADL, they have to replace both the O2 stores and metabolize the
lactic acid that has built up

genes and (the development of) behavior

what are the three conditions for natural selection to happen?
1. variation
2. relationship to fitness
3. genetic basis
how can you demonstrate this for behavior? what are the implications of this assumption? how
do genes and environment/experience determine behavior (and other phenotypes?)

genes and behavior

• genotype + environment  behavior

genotype: the genetic material (dna sequence) that the individual inherited from their parent
phenotype: the observable traits (including behavior) of the individual
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nature via nurture

• is behavior innate vs. learned?
• it’s not black and white
virtually all traits (including behavioral traits) develop through a complex genetic-
developmental program – they depend on both genes and experience
• genotype ‘assumes’ and requires a species-typical enviornment to produce species-
typical behavior
 nature via nurture rather than nature vs. nurture

relationship between genotype and phenotype

a given genotype can produce different phenotypes contingent on key environmental differences
(facultative traits or polyphenisms)
• different phenotypes do not necessarily imply different genotypes

evidence for genetic component of behavior

direct evidence:
1. breeding experiments and pedigree studies
2. selection experiments a. directional: against one end; artificial selection b. disruptive:
against the mean c. stabilizing: against the extremes
3. hybrid studies: intermediate results when two species or populations are crossed

breeding experiments
independent and satellites in ruffs
alternative mating strategies
independent: fight for position on lek, https://www.scilifelab.se/news/a-
display for females that arrive supergene-underlies-genetic-differences-in-
testosteron-levels-and-sexual-behaviour-in-
satellites: show up on lek, are not territorial
male-ruff/
or aggressive, and (for some reason) are
tolerated by independents
• AA or Aa → satellite the satellite version of the supergene is
• aa → independent dominant to the independent yet most
males, ~85%, in the population are
the locus involved is in fact a “supergene”
independent — why?
consisting of ~90 genes

a bird with four “sexes”?

independents: (most males > 80%) fight for position on lek, display for females that arrive
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satellites: (~16% of males) show up on lek, are not territorial or aggressive, and (for some
reason) are tolerated by independents
female mimic (faeder): rare morph (< 1% of males) look like female (smaller in size to other
males), have large testes for their size!
both alternative phenotypes are linked to a “super gene” consisting of ~90 genes. (due to an
inversion in chromosome 11, coming about at about 3.8m years ago): homozygous individuals
for these inversion die (explains why faeder are rare) then there was a recombination about
500k years ago, which led to the satellite phenotype

another bird with four “sexes”

(tuttle et al. 2016 curr. bio.) white-throated sparrows have two color morphs: white and tan
white males are more aggressive, promiscuous, not much of a parental care compared to tan
males — white females are also more aggressive and solicit more copulations than tan females

selection experiments
a. directional: against one end (artificial selection a good example)
b. disruptive: against the mean
c. stabilizing: against the extremes

selection for exploratory behavior

almost all behavioral traits that have been tested respond to artificial selection (i.e., almost all
behavioral traits have a heritable basis)

russian silver foxes

 (dmitry belyaev) selection for one trait can produce correlated effects on other traits
 natural selection can move quickly (artificial selection even more quickly)

artificial selection: dog breeds

select for behavior as well as for physical traits

 border collie: herding and not chasing

 water rescue dog

hybrid studies
migration in blackcap warblers

 s. german blackcaps fly soutwest to spain, austrian blackcaps fly southeast to turkey
 what would the offspring of s. german X austrian blackcaps do?
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1. bring adults of the two blackcap populations (from austria and southern germany) into
the lab
2. breed (cross) males and females of the two populations
3. the next autumn, test the preferred migratory direction of the adults (who have migrated
before) and of their naïve hybrid offspring (who have not)
methods: migratory restlessness & emlen results:
funnels

investigating the genetic basis of behavior via developmental studies

the isolation experiment: raise organisms in isolation from the relevant stimuli (e.g. conspecifics,
nesting materials, etc.) or raise them in an altered environment

do the bats have an innate reference for speed of sound? (emichai and yovel, 2021)
bats navigate by echolocation: estimate the distances of objects by the timig of the return of their
calls or clicks
technically, this calculation requires that the abts “know” the speed of sound — is that
something they learn or is that something they “know innately”?
raise pups in normal air vs. helium enriched air where sound travels 15% faster — thus, a
reflected sound from target will arrive sooner, which should make the target appear closer to
the bat

 all bats (whether they were raised in normal or helium enriched air) behave the same
way behave as if the target is closer in heliox!

umwelt: (literally translated as “the world around”) the notion that different species (and
sometimes members of the same species) occupying the same space will have very different
perceptual environment
important to keep in mind when thinking about abimals (very important concept in animal
communication!)

 also watch the documentary “life in color” on netflix

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…but its not always so simple

three examples:
1. imprinting (‘discovered’ by lorenz)
2. white-crown song learning
3. indigo bunting star compass learning
lorenz: imprinting as a model for “constrained” learning
imprinting is common in presocial animals
animals can be imprinted on almost anything they encounter during the sensitive period, and
visual movement is the key

imprinting in salmon
https://www.nps.gov/articles/coho-monitoring-to-understand-change.htm,
• very few salmon make it to the open water (<2% of juveniles)
• of those who make it to open water are able to return, 95% return to the natal stream to
breed  homing to the natal stream!
each stream have a unique chemical composition

 hypothesis: they imprint on this chemical composition (i.e. smell) of their natal stream?

take juvenile salmon, and raise them in the tank containing either morpholine or pea (two
artificial chemicals) mark and release them into the lake michigan
a year later they marked two streams with small amounts of either morpholine or PEA
monitor a bunch of streams (along with the marked stream) to see where the salmon return
most returning smolts end up returning to the stream which contains the artificial chemical (m
or pea) > clear evidence of chemical imprinting

chemical imprinting is not the whole story  they have magnetic imprinting as well

development of behavior
genotype + environment → phenotype
this formula suggests that the G + E process is a simple additive one — but in general, the
process is a much more complex, interactive one
examples:
1. white-crowned sparrow song learning
2. orientation of migration in indigo buntings
what information is encoded in genes? what information is extracted from the environment?
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white-crowned sparrow song learning (tape tutor method)

 take baby bird from nest at 3-4 days old (blind & deaf), hand-raise it to independence
 song-tutor the bird during his natal summer
 record song he sings as adult next spring

white-crowned sparrows have dialects

(marler suspected these dialects were learned)
• marin dialect
• berkeley dialect
• sunset beach dialect

indigo bunting migration

• breed mostly in eastern US
• overwinter in central america
• how do they orient themselves?

steve emlen’s star compass studies

experiment 1:
do buntings use the stars to orient themselves? put adult birds in ‘emlen’ funnels outside
• clear? – they orient south
• cloudy? – they orient randomly
use star compass/map
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experiment 2:
is the “star map” learned? take young birds from nest and hand-raise them

 ½ see sky jul/aug in sept, orient south

 ½ see no sky in sept, orient randomly
yes, birds learn to use the star map! but how?
experiment 3:
hypothesis: young bird learns north star by watching movement of stars, locating stationary
star, & memorizing star patterns around it

a young indigo bird’s genetic code and developmental machinery somehow contains a
“program” to
1. locate the stationary star in the sky
2. learn the star patterns around it (= the “star map”)
3. when conditions are right, head off 180 degrees away from N.
an example of a genetic-developmental program (GDP)
why such a developmental program?

 polaris was not and will not always the “north star” due to the precession of earth — the
precession has a period of 26000 years: approximately 13000 years ago and 13000 from
now, the north star will be vega in the constellation lyra
 13000 years is a trivial amount of time in evolution of a species (songbirds evolved
about 50 million years ago)
perhaps it is better to learn “fixed star” rule than thhe star map itself!

why be social?
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altruism: behavior that benefits another individual at a cost to the altruit’s personal (direct)
fitness (ability to produce offspring)
–C +B

how could altruism evolve?

is a seemingly altruistic behavior really altruism?

 not really altruism

 simple cooperation or mutualism (+,+)
 true altruism (–,+)
 kin selection
 reciprocal altruism

example:
• feeding of penguins in a crèche (not really altruism)
• cooperative hunting in wolves (+, +)

altruism & kin selection

kin selection: selection of a trait through helping relatives, either
1. descendant kin (offspring) — direct selection
2. non-descendant (collateral) relatives — indirect selection
inclusive fitness = direct + indirect fitness
(usually, when people talk about kin selection, they mean indirect selection)

three factors are important in the spread and maintenance of an altruism gene by kin selection:
1. benefit to recipient, B
2. cost to altruist, C
3. degree of relatedness between altruist and recipient, r
coefficient of relatedness, r
r = the proportion of genes identical by descent (IBD

hamilton’s rule states the conditions under which altruism will spread — in it simplest form, it
is:
rB > C
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Σ (rBs) > C there can be multiple beneficiaries

j. b. s. haldane: m
when should you be altruistic?
when B > C/r
Recipient

more general form of hamilton’s rule

rB B > rC C
rB = r of actor to the recipient
rC = r of actor to individual suffering cost
for example:
help sibling raise his/her kids (your nieces/nephews)

altruism and kind selection

kin selection
an example: american turkeys
related males frequently form displaying pairs to attract females (females prefer duos to single)
but: only the dominant turkey mates, i.e., subordinate turkey is an altruist

hypothesis: the dominant/subordinate duos are btrother and because females prefer duos to
singles, the subordinate male gets more through kin selection then he would going ın his own,
i.e. this altruistic behavior of the subordinate

altruism and reciprocity

reciprocity: altruist repaid at a later date by the recipient

you other
time 1 –C +B
time 2 +B –C
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time n … …
average B–C B–C

prisoner’s dilemma
prisoner a can not implicate prisoner b (‘cooperate’) or squeal om him (‘defect’) and ditto for
prisoner b re A if both cooperate, do better than if both defect. what should they do?

player b

cooperate defect

cooperate reward for mutual maximum

cooperation (only 1 punishment (10
player a year in prison) years in prison)

defect maximum reward punishment for

(freedom = 0 years in mutual defection (5
prison) years in prison)

player b

cooperate defect

cooperate B–C

player a
defect maximum reward punishment for
(freedom = 0 years in mutual defection (5
prison) years in prison)

player b

cooperate defect

cooperate reward for mutual maximum

cooperation (only 1 punishment (10
player a year in prison) years in prison)

defect maximum reward punishment for

(freedom = 0 years in mutual defection (5
prison) years in prison)
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but these conditions hold only in a blind, one-shot game

situation is changed if we change any of the implied assumptions:
1. repeated interactions with individuals
2. recognition/memory: remember who cheats and who reciprocates
3. flexible behavior: can modify behavior (give or not give) depending on the past
interactions

axelrod and hamilton (1981) (computer tournament)

“tit-for-tat” is an ESS
(1) cooperate on 1st play
(2) do what your partner did on the previous play

reciprocal altruism in vampire bats (desmodus rotundus)

hungry, unsuccessful bats beg for food from non-related partners

vampire bats appear to meet the conditions for tit-fot-tat ESS

1. long-term associations, therefore opportunity for repeated interactions with roles
reserved
2. probably individual recognition (olfactory)
3. probably remember (in lab, more likely to regurgitate to roost mates than to strangers)

example of reciprocals altruism

hamlets (a kind of fish): simultaneous hermaphrodites
which role should you take?
♀ or ♂
what they do:
• mate 1x/day
• parceling
• “egg trading”

what drives sexual selection?

trivers: key to sexual selection is parental investment (PI) = effort to put into your
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eusociality
s

altruism, helpers at nest and den

why help?
white-fronted bee-eaters
helping explained mainly by kin selection in bee-eaters

animal communication
animal communication
1. communication basics
2. types of communication signals
3. the problem of honest signals
4. phylogeny of signals

communication – basics
communication:

 transmission of information via a signal benefiting both signaler & receiver (on average,
over evolutionary history, in most contexts)
information:

 reduction of uncertainty

signal:

 adapted for signal function

why/what do animals need to signal? animals are uncertain about states of the world, they need
information
the 3 F’s (food, fight, fuck) + P (predator) + I (identity)
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• where’s the food?

• do you wanna fight? (dominance)
• do you wanna have sex?
• what/where is the predator?
• who are you? (identity)
animals can ‘communicate’ without signals — they have to make decisions, and so they use
whatever information is available
• prior probabilities regarding animals in general
• prior probabilities regarding that animal in particular
• cues that animal gives
• signals that animal gives

cues vs. signals

cues are “leaked”:

 easily-assessed information that the animal may not ‘want’ to reveal

signals are functional:

 contain information that animal ‘wants’ to reveal, because it benefits it

(benefit always measured relative to not signaling)

• does animal need to signal (i.e., what are the benefits)?

• is the behavior/action a signal or merely a cue?
• if it is a signal, what is the evidence of it having been elaborated for signal function?
• is it an honest signal (does it convey true information)?

types of communication signals

1. information about external world
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a. food signals
b. predator (warning/alarm) signals
2. information about identity
a. interspecific
(1) warning
(2) mutualism
b. intraspecific
(1) species
(2) individual
(3) group
(4) kinship
(5) status
3. signals that indicate motivational state and/or synchronize social interactions
a. threat
b. reassurance
c. submission/appeasement
d. courtship & mating
(1) epigamic aspects (mate attraction)
(2) motivational/timing aspects
e. parent-offspring communication
f. intra-group (cohesion) signals
g. inter-group (spacing) signals
4. assessment [overlaps with all else]
a. mate attraction context
b. threat context
c. predation context

information about external world: food signals

• bee dance: https://www.youtube.com/watch?v=-7ijI-g4jHg

• ravens: “food bonanza here!”
• chimps
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information about external world: predator signals

belding’s ground squirrel: ground predator (coyote) & aerial predator (hawk) calls
three kinds of alarm calls (for eagles, leopards, and snakes) in vervet monkeys:
https://www.youtube.com/watch?v=3lsF83rHKFc

white-tailed deer: “upturned tail” gesture — the intended receiver is the predator (“i saw you”)

information about identity: interspecific — warning

a mullerian mimic
mullerian mimicry in poison dart frogs
the r. imitator morph changes depending on
what the other common poison dart frog is
in the area!

signal sharing: (black & yellow stripes)

• yellowjacket
• masarid wasp (a mullerian mimic)
• a beetle (a batesian mimic)
• a fly (a batesian mimic)
coral snake (poisonous) — mountain king snake (non-poisonous)
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 predators stay away from mountain king snake as well

sea slugs (a variety of complex patterns and vibrant colors)

 basically, possible predators stay away from all of these “weird-looking” things (warning
coloration or neophobia)
mullerian mimicry in sea slugs:
• 4 different species of genus chromodoris

information about identity: interspecific — mutualism

cleaner fish tend to have a blue and yellow stripes more frequently than related fish that are
non-cleaner fish
• blue gives the most contrast in front of reef
• yellow gives the most contrast in front of blue water
cleaner fish and its ‘clients’: https://www.youtube.com/watch?v=V5EszU8yuA
an aggressive mimic of the cleaner-wrasse: sabre-toothed blenny

information about identity: intraspecific — species

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tree frog

information about identity: intraspecific — individual

marsh wren

information about identity: intraspecific — group

budgerigars

information about identity: intraspecific — kinship

praire dogs
sometimes you may expect this kind of signal not to evolve (in favor of the offspring of extra-
pair copulations)

information about identity: intraspecific — status

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status signaling — harris’ sparrows

dynamic signals that indicate motivational state and/or synchronize social interactions: threat

a dog

anolis lizard

frilled lizard

hooded seal:
http://arkive.org/hooded-seal/cystophora-
cristata/video-12.html
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yellow baboon open-mouth threat

chimp tense-mouth threat

vervet monkey — red, white & blue display

. . . and other unique threat signals

dynamic signals that indicate motivational state and/or synchronize social interactions:
reassurance
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dynamic signals that indicate motivational state and/or synchronize social interactions:
submission/appeasement

the difference between active vs. passive

submission signals:
who initiates the signal

dynamic signals that indicate motivational state and/or synchronize social interactions:
epigamic aspects (mate attraction)

mandarin duck
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goldeneye duck
animals often use ‘songs’ to attract mates from a distance
• tree frog
• marsh wren
• blue whale (and so on)
vibratory signals — male attraction

jumping spider courtship

dynamic signals that indicate motivational state and/or synchronize social interactions:
motivational/timing aspects

whooping cranes
george archibald breaking the code!
• http://www.youtube.com/watch?v=rh6FJpBjQOA
just four days old chick is fed with a whooping crane hand puppet:
• http://www.youtube.com/watch?v=JzQd-ZrTwJU&feature=youtu.be
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olfactory estrus signals: synchronizing

dynamic signals that indicate motivational state and/or synchronize social interactions: parent-
offspring communication

dynamic signals that indicate motivational state and/or synchronize social interactions: intra-
group (cohesion) signals

dynamic signals that indicate motivational state and/or synchronize social interactions: inter-
group (spacing) signals
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marsh wren

howler monkey

cheetah

assessment: mate attraction context

epigamic signals often used to assess the signaler (mate attraction context)

peacock spider
http://www.youtube.com/watch?v=9GgAbyYDFeg

assessment: threat context

springboks pronking/stotting
http://www.youtube.com/watch?v=8Ba3UxqXiXU
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review: types of communication signals

1. information about external world
c. food signals
d. predator (warning/alarm) signals
2. information about identity
c. interspecific
(1) warning
(2) mutualism
d. intraspecific
(1) species
(2) individual
(3) group
(4) kinship
(5) status
3. signals that indicate motivational state and/or synchronize social interactions
h. threat
i. reassurance
j. submission/appeasement
k. courtship & mating
(1) epigamic aspects (mate attraction)
(2) motivational/timing aspects
l. parent-offspring communication
m. intra-group (cohesion) signals
n. inter-group (spacing) signals
4. assessment [overlaps with all else]
d. mate attraction context
e. threat context
f. predation context

the problem of honest signals

communication: transmission of information via a signal benefiting both signaler & receiver (on
average, over evolutionary history, in most contexts)
sender

+ –
+ “true” eaves-
receiver communicatio dropping
n
– deceit or
manipulation
some examples of deceit (sender +, receiver –) or eavesdropping (sender –, receiver +) when
sender and receiver are different species
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interspecific communication

an example of deception — but not really deceptive communication

a mimic — deception

eavesdropping

deception — femme fatales

sender
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+ –
+ “true” eaves-
receiver communicatio dropping
n
– deceit or
manipulation

some people argue that signaling often does not benefit both parties simultaneously, giving
examples of aggressive eavesdropping or deceit/manipulation
but clearly, these cases are possible only because they parasitize honest (‘true’) communication
femme fatale fireflies

 their strategy works only because other species honestly signal their species
frog-eating bats

 their strategy only works because frogs must advertise their species

batesian mimics

 works only because warning colors are usually honest

how is honesty maintained in intraspecific signaling?

make the signal
• costly,
• constrain it,
• or police it
we’ll consider the following examples:

 wolf dominance — subordinance signals (policing)

 harris’ sparrows — status signals (policing and/or cost)
 european toads — “deep croak” hypothesis (constraint)
 red deer — aggressive displays (cost/constraint)
 tungara frog — mate attraction call (cost)

policing
the simplest case
in long-term social groups with individual recognition—like wolves—honest signaling can be
maintained by policing — that is, bluffs usually are called
in fighting context, dishonesty is often not worth the risk
it can pay to be honest: even the stronger party could get hurt, and the weaker party can live to
win another day
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status signaling in harris’ sparrows

dye blacker: birds do not rise in hierarchy

 they are challenged frequently, back down, move to edge or out of flock

dye whiter: birds are challenged much more,

 don’t back down, become ‘paranoid’

need to look and feel dominant (or subordinate)

then why not cheat? well, how hard would it be to cheat? all you need is black head/bib feathers
plus a high level of testosterone!
• it may be costly for poor-condition birds to maintain high level of testosterone
• policing ensures that stronger birds will defeat weaker birds so cheating does not pay,
honest signals save time

how is honesty maintained?

constraint + assessment
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european toads — “deep croak” theory: because of physical constraint, lowpitched croaks
honestly reflect size and competitive ability (index signal)
roaring contests in red deer stags — how loud and long a stag can roar is limited (constrained)
by his condition, endurance; thus the intensity of this signal is an ‘index’ to his fighting ability
signaling system is hierarchichal

handicap model of signal honesty

cost or constraint ensures honesty

costly signaling
tungara frog (physalaemus pustulosus) adds
‘chuck’ to end of its advertisement calls
(whine-chuck)
experiments show that females prefer a call
with a chuck
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females prefer a call with the chuck — why? does chuck make the call more costly? yes!

frog-eating bats can more easily localize frogs that add the chuck!

sensory exploitation
manipulation theories put emphasis on benefit to sender and posit that responding to the signal
does not have to benefit the receiver
sensory exploitation theory is the best example of this kind of theory
sensory exploitation theory: novel or extreme traits attract receiver by triggering a pre-existing
sensory bias
should we always assume receiver benefits by noticing the signal? — perhaps receiver simply
can’t help it!
finches: females prefer males with artificial feather hats over normal males!
is this (inherently) honest signaling?

swordtails have a sword, but closely related platyfish do not

 experiments show that females of both species prefer males with swords!

cichlid fish, bird of paradise, and so on..

• true communication? or sales pitch / propaganda? or simple manipulation?
we return to this issue in the next section sexual selection and mate choice
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phylogeny of signals
the elaboration of actions/traits for signal function
ritualization: the evolutionary process whereby a behavior is elaborated to become a more
effective communication signal
any trait—whether behavioral, physiological, or morphological—can be ritualized for
communication
intention movements:
• showing teeth (“do not come closer or i will attack”)
• opening wings wide open (“i am about to fly”)
panda principle:
evolution acts by modifying existing structures (traits) rather by designing them from scratch
• therefore although the trait will be adaptive, its ‘design’ will often appear to be non-
optimal
“imperfections are the primary proofs that evolution has occurred, since optimal designs erase
all signposts of history” (s. j. gould)
a nice example of ‘panda principle’ — balloon fly (covered earlier in the course)

panda’s ‘thumb’ is a modified wrist bone (it

is an evolutionary contraption or
contrivance)
the radial sesamoid was co-opted to serve
its present role in handling bamboo
“it is jury-rigged, it is a rube goldberg
device”

reproductive strategies, sexual selection

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reproductive strategies
a set of behavioral and physical adaptations designed specifically to maximize an individual’s
mating and reproductive success

topics:
1. why sex?
2. males vs. female
3. sexual selection

why NOT sex?

it is costly!
sex vs. asexual reproduction (cloning)

 gonads are expensive organs to produce and maintain

 mating is risky, costly, time-consuming, often involving elaborate structures and
behaviors
 cost of meiosis – this is perhaps the most significant cost
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since each sexually-produced offspring only contains half the genetic material of each parent,
there is a 50% reduction in fitness compared to asexual reproduction

why sex?
benefits of sexual reproduction
everything changes: your competitors evolve, the environmental circumstances change etc.
• evolution is a constant ‘arms race,’ so to speak
genetic variability among offspring permits response to changing biotic environment
red queen hypothesis:
whereas asexual reproduction is a ‘sitting duck’ target for competitors, pathogens, predators, sex
and genetic recombination present a moving target

how to define males and females?

♀ ♂
sex chromosomes are not generally a useful criterion to define males and females
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 mammals

• XY – male
• XX – female

 birds
• XX – male
• XZ – female

 hymenopterans:

• N – male
• 2N – female

 hermaphroditic species

 some lizards, turtles, alligators

• facultative (incubation is temperature dependent)

instead,
‘male’ and ‘female’ are defined by relative size of their gametes (sex cells):

 females produce relatively few but expensive (provisioned) sex cells

 males produce many more but cheap sex cells

anisogamy, i.e., “unequal gametes”

• eggs: few, large, energetically expensive
• sperm: many, small, cheap

sexual selection
usually trades off with natural selection, i.e., traits that increase mating success usually decrease
survival and/or parental investment
occurs when
(1) males and females compete among themselves for access to mates and resources
associated with breeding (intrasexual selection)
(2) and/or males and females try to attract females or males by flashy colors, behaviors etc.
(epigamic selection)
so there are two types of sexual selection
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intrasexual selection
male-to-male or female-to-female competition

generally involves  weapons & fighting

• https://www.youtube.com/watch?v=GDiurxZtmxQ
• https://www.youtube.com/watch?v=-VWFreC4onI
• https://www.youtube.com/watch?v=ULRtdk-3Yh4

epigamic sexual selection

males try to attract females (or vice versa, in rarer cases) by flashy colors, behaviors etc.

note that this northern jacana is not a male!

(sex-role reversal)
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flip side of epigamic selection: female choice

what exactly are females choosing?
1. https://www.youtube.com/watch?v=1XkPeN3AWIE

reproductive strategies
1. a set of behavioral and physical adaptations designed specifically to maximize an
individual’s mating and reproductive success
2. a set of behavioral and physical adaptations shaped by sexual selection

sexual selection tends to produce

• weapons
• and elaborate ornaments and displays

what drives sexual selection?

trivers: key to sexual selection is parental investment (PI) = effort put into helping your
offspring reach reproductive age
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parental investment (PI)

initial PI
females produce large egg cells, males produce small sperm
• female's provisioning of egg cell
specialized PI
specialized adaptations (mammary glands, incubation patches, etc.) directly related to
reproduction (e.g., gestation, nursing)
usually supplied by one sex only, and that is almost always the female
• female’s nourishing of the embryo
• female’s producing yolk-filled egg (reptiles and birds)
• incubation (reptiles and birds)
• female lactation (in mammals)
generalized PI
generalized/behavioral adaptations (and costs)
usually behavioral and can be supplied by either sex, more commonly the female in mammals
• guarding the offspring
• feeding the offspring
• defending a territory/nest used for feeding and protecting young

in general,

 female PI is much greater than male PI

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parental investment theory

trivers: key to sexual selection is parental investment (PI)
sexual selection:
1. sex investing less will compete for sex investing more (intrasexual selection)
2. sex investing more will be discriminating in choosing mates (epigamic selection)
typically: female PI > male PI
so, usually,
• its males competing among themselves for females
• and females choosing among males

• females as the discriminating sex and males as the less discriminating sex
an important concept: reproductive variance (RV)
• satin bowerbird as an example
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trivers: key to sexual selection is parental investment (PI)

typically:

♀ PI > ♂ PI (i.e., ♂ RV > ♀ RV) so usually ♂ ♂ compete among themselves for ♀ ♀ and ♀ ♀
are choosing among ♂ ♂

that is

• female PI > male PI  male RV > female RV

• males compete, females choose
or

• when female PI  male PI  female RV  male RV

• competition more equal, less intense
how do we test trivers’ hypothesis that PI drives sexual selection?
look for cases of “sex role reversal”!

• male PI > female PI  female RV > male RV

• females compete, males choose

sex role reversals

• seahorses & pipefish — specialized PI, male ‘gestation’ & lactation
• mormon crickets — spermatophores
• jacana — polyandry, ♂ parental care)

in all these cases, males are the more choosey sex and/or females compete among themselves
for males
trivers would explain these cases so:

 males invest more than females, so females will compete over males and males will be
the more choosey sex
however,

 why is male PI > female PI in these sex-role reversed species in the first place?
it may be that PI is driven by sexual selection and not the other way round!

reproductive strategies, mate choice

mate choice
begin with simplest, most common case:
female is choosing male
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cases of pure epigamic selection

red-capped manakins’ moonwalk display:
• http://www.youtube.com/watch?v=PDYpW3zyXqQ
• http://www.youtube.com/watch?v=kkSxRRxvQ8E
• http://www.youtube.com/watch?v=eI_quJRRGxk
blue manakins’ communal display:
• http://thekidshouldseethis.com/post/male-blue-manakins-wait-in-line-to-impress-a-
female

vs.

mate choice and sexual selection

direct benefits: female choice leads to more resources (hence more offspring)
good genes: female choice leads to improved genetic quality of offspring in respect to survival
runaway selection: female preference increases because it is linked to ‘sexy son’ advantage
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sensory exploitation: male evolves display trait that exploits pre-existing sensory bias in female
chase-away selection: female preference then evolves away from male traits if there is a cost to
being exploited
note: these are not mutually exclusive!

direct benefits
when resources (good nest sites, good food) differ between territories of males, females may
choose purely on the basis of resources (though presumably ‘better’ males would get better
territories)

female may choose on basis of ‘real estate’

what happens if we add 2 females to territory 1?
polygyny threshold model
female may prefer mated male (polygyny) rather than unmated male (monogamy) if she gets
more direct benefits from that choice (resource defense polygyny)

good genes
honest signals are uncheatable, and provide
accurate information about health, vigor,
and general condition
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females choose males with more eyespots…

but is it because they have more eyespots?

 yes

females prefer males with more eyespots...

but are they better quality males?

 yes

males with more eyespots are healthier…

but is this fitness different heritable?

 yes

a study with peafowls 8 males (peacocks) x 4 females (peahens), each 96 offspring released:
• % of chicks surviving after 2 years and mean are of their father’s eyespots (mm2) is
positively correlated!
• apparently, they are better quality males (pass on better genes)
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honest signaling: handicap model

are peacock’s gorgeous feathers indicators of good genes because they are handicaps?
an epigamic signal will be an honest indicator of the condition (quality) of the male if it is less
costly (a smaller handicap) for good condition (high quality) males than it is for poor condition
(low quality) males

indicator trait (ornament or display) is a handicap in that it is costly or difficult to produce, and
is supported by many genes affecting overall fitness, such as genes for disease resistance —
contrast this with runaway processes

fisher: runaway selection

are peacock’s gorgeous feathers the result of a runaway process?
https://www.youtube.com/watch?v=koSvAh6za7Y

lengthened (‘supernormal’) tails increase

male’s mating success (mean number of
nests per male)!

the basic idea is that female preference for a male trait (say tail length) will be favored because
female with that preference will have ‘sexy’ successful sons (thus increasing her RS) (& this
assumes genetic basis of preferance)
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best test: we should find a genetic correlation between genes for female preference and genes
for male trait
“runaway” refers to the positive feedback between trait and preference — genes for trait and
preference will become associated

bakker (1993): sticklebacks

took fish from stickleback populations that
varied in terms of brightness of male nuptial
coloration (probably related to intensity of
predation)
crossed fish from bright population with fish
from dull population
sons’ intensity of red correlated with that of
their fathers’ and with the daughters’
preference for red!

where does it stop?

when benefit (in respect to mating advantage) is maximally different from its cost (in respect to
survival)

sensory exploitation
are peacock’s gorgeous feathers have their origins in
sensory exploitation?
sensory exploitation: male evolves display trait that
exploits pre-existing sensory bias in female
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these hypoteses are not mutually exclusive!

direct benefits: female choice leads to more resources (hence more offspring)

 female might choose male for direct benefits and good genes, or she might pick different
males for each
good genes: female choice leads to improved genetic quality of offspring in respect to survival

 needs to be demonstrated, often just assumed to be true!

runaway selection: female preference increases because it is linked to ‘sexy son’ advantage

 doesn’t say anything about why female has preference, just that the preference itself can
be a driving force
sensory exploitation: male evolves display trait that exploits pre-existing sensory bias in female

 predicts the kind of trait (one for which she has a sensory bias) female might choose in
first place
chase-away selection: female preference then evolves away from male traits if there is a cost to
being exploited

 assumes sensory exploitation as pre-condition, and that there is a cost to female of being
exploited

sexual selection is directional selection: it can drive epigamic trait up or down

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in evolution you should expect not only to

find cases where elaboration of epigamic
trait has increased (through female choice)
but cases where it is has decreased (through
predation and other opposing pressures)

summary
assuming that female is the choosing sex:
1. in some cases, female chooses for direct benefits (not male epigamic traits)
2. when female gets only genes, she should be selected to choose good genes, but this
requires reliable indicator traits
3. reliable indicator traits will usually be costly ones, thus only best males develop best
signals (handicap hypothesis)
4. female choice can lead to superexaggerated traits (runaway selection)
5. runaway selection will be stopped when costs of trait for males exceed its benefits
6. the kind of male trait that is selected may be one for which the female already has a
sensory bias (sensory exploitation)
7. females should be selected to show sales resistance to signals that cost them (chase-
away selection)

mating systems
key concepts of mating systems
1. mating system = mating and parental care
2. monogamy, polygyny, polyandry
a. bonding definition
b. reproductive variance definition
3. reproductive effort
a. ME (mating effort) & PE (parental effort) usually mutually exclusive
b. but sometimes not! (sticklebacks)
4. other interesting cases
a. rhea
b. pipefish & seahorses (phylogeny!)
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5. intensity of sexual selection

a. often reflects apportionment of ME and PE
b. tricky case: both sexes bright!
6. human mating systems

dimensions of mating systems

bonding:
do male and female have an arrangement that goes beyond copulation?
multiple mating:
no? if yes: by male only? by female only? by both? if both, which sex mates with more
individuals?
parental care:
both? female only? male only? neither?

mating systems terminology derived from anthropology

monogamy
• ♀ ♂
polygyny

• ♀♀♂
polyandry

• ♀♂♂

mating systems in terms of reproducive variance

reproductive variance (RV): variance in reproductive systems

 important point: the degree of asymmetry in RV is an indicator of the intensity of sexual

selection
polygyny: male RV > female RV
monogamy: male RV = female RV
polyandry: female RV > male RV
polygamy: polygyny or polyandry
promiscuity: no bonds, males and females mate multiple times

other component of the mating system is the form of parental care

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usually:

 polygyny – female parental care

 monogamy – biparental care
 polyandry – male parental care

for example, an extreme case: elephant seals

https://en.wikipedia.org/wiki/northern_elephant_seal
• polygyny
• female parental care
• male RV > female RV

primate mating systems

polygynous ones:
• gorillas — single-male groups (harems)
• chimpanzees — multi-male groups
monogamous ones:
• siamangs
• owl monkeys
monogamy & shared parental care:
gibbons & siamangs

 territorial
 young dependent on parents until age 3
 parental care primarily by mother in the 1st year, but switches to father in the 2nd year
cotton-top tamarins
monogamy occurs in groups where a sufficient number of older offspring remain to help
polyandry occurs in groups lacking old offspring — in these situations, pairs recruit the help of
additional male

return to trivers: “differential PI drives sexual selection”

female PI

• sexual selection between ♂ – ♂

and if

• ♂ PI > ♀ PI,
then get role reversal
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a problem for trivers’ theory

cases where male PI > female PI but male RV > female RV & no role reversal
this pattern is common in fish:
• male makes nest, female spawns & leaves, male defends the eggs
trivers’s assumptions:
• PI sex difference is prior (cause, not effect)
• PI and mating effort (ME) are mutually exclusive (trade off)

reproductive effort
mating effort + parental effort

or

in this view ME and PE are mutually exclusive

reproductive strategies
1. a set of behavioral and physical adaptations designed specifically to maximize an
individual’s mating and reproductive success
2. a set of behavioral and physical adaptations shaped by sexual selection
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3. a program for the optimal allocation of reproductive effort

what if ME and PE are not mutually exclusive?

stickleback fish
in most fish male PI > female PI but no sex role reversal
in sticklebacks, though male has to care for eggs in the nest, he can still court females and the
presence of eggs attracts additional mates! ME & PI often not mutually exclusive in fish!
• why are males with eggs in the nest more attractive to females?
• why do “gals like guys with dogs”?
rhea
(south america) a bird with male-only parental care but no sex role reversal
• harem polygyny
• male incubation & parental care
• + sequential polyandry!
what factors might favor male parental care in rheas?
which sex loses more by desertion?
which sex is better equipped for parental care?
which sex will take longer to get ready for next round of mating?

concept of operational sex ratio

primary sex ratio = number of males divided by the number of females

 ♂:♀
 PSR = 1:1
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operational sex ratio = number sexually receptive of males divided by the number of sexually
receptive females

 for example, potential reproductive rate of males might be higher than that of females
 ♂♂♂♂♂♂♂♂♂♂:♀
 OSR = 10:1

pipefish & seahorses

males receive the gametes, are pregnant, and give birth
after the female deposits her unfertilized eggs into the male, the outer shell of the eggs breaks
down, and tissue from the male grows up around the eggs in the pouch
after fertilizing the eggs, the male closely controls the prenatal environment of the embryos in
his pouch
the male keeps blood flowing around the embryos and provides oxygen and nutrition to the
developing offspring through a placenta-like structure until he gives birth

 ♂ PI > ♀ PI, sex role reversal

importance of evolutionary history:

steps to male parental care (and sometimes polyandry) in seahorses & pipefish

male parental care of eggs

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then  male specialization for parental care & female ability to lay more eggs than male can care
for

then  female competition to obtain several males (or mate guarding)

intensity of sexual seleciton

red-winged blackbird  polygynous

sexual selection can occur even in

 monogamous species with biparental care – its just less intense

northern cardinal  monogamous

• dimorphic – ♀ somewhat colorful

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house finch  monogamous

• dimorphic – both sexes less colorful than cardinal

bank swallow  monogamous

• monormorphic – differences are behavioral only

eclectus parrots  monogamous

• dimorphic

human mating systems & mate choice

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• monogamy most common

• but sequential, also extra-pair mating
• polygyny (mild) also fairly common
• polyandry (milder) also found (rare)
• bi-parental care, division of labor

animal mind
speculations about animal mind center on one or another of these attributes
intelligence:

 are animals smart, do they reason problems out?

complex cognitive mechanisms:

 do they use complex mental mechanisms such as mental maps?

consciousness:

 what is the content of the animal mind, do they introspect?

emotion:

 do animals have feelings like us: love, hate, jealousy, sadness, etc.?
intentionality:

 do animals make plans?

self-awareness:

 are animals aware of themselves as unique individuals?

theory of mind:

 is an animal aware that other animals have minds too?

communication:

 can animals ‘talk’ with one another? with us?

special problems with studying animal mind that we don’t have when studying animal behavior
1. problem of objectivity: can’t know animal mind directly. behavior is observable, mental
processes are inferred
2. problem of anthropomorphism (the injection of human qualities into animals): humans
are naturally inclined to assume that animals think and feel like we do

anthropomorphism & theory of mind

humans seem naturally inclined to engage in anthropomorphism, maybe because humans
develop a refined theory of mind which they routinely use to interpret others’ behavior
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theory of mind (TOM): the awareness that

others have minds as well as yourself

TOM develops with age — young children act

as if they are unable to distinguish between
what they know and what others know

two separate points about TOM:

1. humans instinctively apply TOM to
animals (a mentalistic, anthropomorphic
approach)

2. do animals too develop TOM? (we deal

with this one later)

historically, two distinct approaches to the study of animal mind

mentalistic approach: anthropomorphic – investigator tries to demonstrate that animal employs
human-like mental processes in dealing with its world
mechanistic approach: investigator assumes that animal is like machine in some respect,
proposes a simple mechanical model of the underlying mental processes

animal mind lecture outline

1. mechanistic and mentalistic approach to animal thinking
2. background: two influential mechanistic viewpoints:
o strict operational behaviorists or S.O.B.s (simple learning)
o classical ethologists (instinctive behavior)

3. some examples:

 indicating apparent limitations of animal thinking

 which seem to require a higher order of thinking
 which may indicate animal awareness/consciousness
 teaching language to animals
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how similar is animal thinking to us?

mechanistic approach mentalistic approach

start from the study of observable behavior
alone
always explain behavior by the simplest
possible mechanism (law of parsimony)
assume animals are like simple machines
until proved otherwise
explain animal minds by reference to mind
we know the best: human mind
hypothesis: animal mental processes are
analogous to human mental processes
assume animals are like humans until proved
otherwise

mechanists complaints with mentalists: mentalists complaint with mechanists:

• lack of parsimony • simple-minded
• lack of objectivity • pig-headed
• anthropomorphism
• anecdotal method
most animal behaviorists, past and present, fall into one of 3 groups:
(1) mentalists
(2) mechanists (including classical ethologists, S.O.B.s)
(3) ‘agnostics’ (most common)

a mechanistic approach: strict operational behaviorists (S.O.B.s)

operant conditioning: behavior is voluntary and goal directed (‘operates on the environment’)
and it is ‘shaped’ by its consequences – strengthened if rewarded, weakened if punished
complex behavior can be ‘shaped’ — the result is a response chain
shaping by successive approximations
(1) https://www.youtube.com/watch?v=ttG7ulOUHtw
(2) https://www.youtube.com/watch?v=TtfQlkGwE2U&feature=related

a mechanistic approach: classical ethology

mechanistic models of instinctive behavior
sign stimulus (releaser)
↓
innate release mechanism
↓
fixed action pattern
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supernormal stimulus

examples from natural or semi-natural situations indicating limitations of animals’ thinking

• chiszar snake experiment
• rothstein robin experiment
• langur ruse (sarah hrdy): deceptive ‘estrus’ signal

chiszar’s snake experiment background:

copperheads prey on both mammals (e.g.,
mice) and amphibians (e.g., frogs)
hunting and feeding strategy
1. locate prey
2. strike and envenomate prey
3. wait 10 minutes and then search for
(now dead) victim

rothstein robin experiment context: some

host species accept cowbird nest parasitism,
some reject
what you need to know about robins (true
of many birds):
1. will eject cracked own eggs or
cowbird eggs;
2. will desert after whole clutch loss or
multiple egg loss
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robins do not immediately leave after the

majority of their eggs were replaced with of
cowbirds’, rather, they first remove the alien
eggs, and then leave

results:

some examples which seem to require a higher

order of thinking
1. insight learning
2. tool using
3. cultural transmission
4. self-awareness (consciousness)
5. language (human  animal)

insight learning
suddenness of finding solution thought to
indicate mental process of ‘insight’

kohler’s chimps

skinnerians components that go into the solution to the

problem
do not assume that chimps (or humans) are
smarter than ‘simpler’ animals like a rat or a
pigeon!
rat or pigeon may just lack the necessary
background experience or the behavioral
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S.O.B.’s have a different take
epstein’s pigeon ‘simulation’ experiment:
 teach pigeon to climb on box and
peck at toy banana
 separately, teach pigeon to push box
to target (spot on floor)
 test day: banana out of reach, box
present – what does pigeon do?
once the pigeon has learned the separate
components – to push the box to a target
and to peck at the banana – it will put them
together quickly and suddenly (insight!) to
get at the banana when it is out of reach

tool use
in chimps
discovered by jane goodall in 1960, 1st known example of tool making and usage outside of
humans, causing louis leakey to declare “now we must redefine tool, redefine man, or accept
chimpanzees as humans”
in gorillas
leah first looked at the new elephant pool and the branch she later used as the walking stick, and
entered the water without the tool — after re-entering the pool and taking the branch with her
right hand, she walked bipedally 8–10 m into the water, frequently testing water depth
sea otters and their favorite rocks
• http://www.youtube.com/watch?v=MdRlD35rl3g&NR=1
does tool use necessarily imply a higher order of thinking? maybe it’s just a specific adaptation…

‘woodpecker’ finch of galapagos islands

woodpecker finch fills niche occupied by
woodpeckers on the mainland (south
america)

new caledonian crow — similarly, fills niche occupied by other species on the mainlands
(australia, new zealand)
http://www.youtube.com/watch?v=TtmLVP0HvDg&feature=related

striped possum (dactylopsila trivirgata) fills “woodpecker” niche in new guinea and australia (where
it is being replaced by related sugar glider)

• nocturnal and arboreal and uses prehensile tail, powerful incisors and elongated 4 th finger to
take beetle larvae and caterpillars from tree bark
• it detects the larvae from inside the trees by rapidly drumming on them with the toes of its
forefoot
• the fourth finger has an unusual hooked nail which it uses to extract insects out of cracks
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cultural transmission
• fongoli chimps (‘ape genius’ movie)
• potato washing in japanese macaques of koshima island

cultural transmission generally assumed to require

(1) insight learning and
(2) observational learning or teaching
caution – has the observing bird learned how to open the bottle or simply that there’s food in it?

some examples which may indicate animal self-awareness/consciousness

the mirror test & self-awareness
gallup and others have argued that behavior directed towards bodily marks visible only in a
mirror may serve as an operational definition of “self-awareness”
using this definition, gallup claimed that only humans, chimpanzees, orangutans, and perhaps
gorillas may be labeled “self-aware”
since then, more animals have passed the test — and quite a few have failed it
gallup‘s procedure:
while animal is anesthetized, the upper brow and ear are smeared with rouge
upon awakening, the animal is exposed to a mirror
if the animal investigates his or her body and touches the rougemarked location, such behavior
is interpreted as evidence of “self-awareness”
http://www.youtube.com/watch?v=vJFo3trMuD8
is self recognition uniquely human?
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epstein’s simulation of the mirror test

epstein, lanza & skinner (science 1981): mirror experiment with pigeons

train pigeon:
a. to localize the origin of blue stimuli briefly reflected in a mirror, and
b. in absence of mirror, to peck at stick-on dots pasted to the feathers they can see
(a) familiarizes bird with properties of a mirror, while (b) establishes a history of self-directed
behavior under ‘no mirror’ conditions
test day: blue dot placed on the pigeons' breast feathers but covered with a cotton bib
once the mirror is uncovered, however, the dot under the bib is visible in the mirror—if—bird
stands upright with its beak pointed toward the mirror
• https://www.youtube.com/watch?v=0Ga5o9cyg9A
• http://www.youtube.com/watch?v=HRVGA9zxXzk

language: what separates humans from other animals?

other ‘unique’ human abilities have been found in other animals
• tool use
• cooperation
• cultural transmission
• use of symbols
• theory of mind
language has been taught to:
• orangutans • dolphins
• gorillas • sea lions
• chimpanzees • parrots
• bonobos • dogs

language learning in animals

original goal: can animals learn language? how big is the gulf between human and animal?
kelloggs raised a chimpanzee as a member of their family — the chimp learned to recognize
more than 100 words, but chimps are not able to produce human speech sounds
sign language
• chimp – washoe (gardners, fouts)
• chimp - nim chimpsky (h. terrace)
• gorilla - koko (p. patterson)
lexigrams
• chimp - sarah, others (premacks)
• bonobo - lana, kanzi (rumbaughs)
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comprehension only
• dolphins (herman)
• calif sea lion (shusterman)
speech
• african grey parrot – alex (irene pepperberg)

lana the chimp

born October 7, 1970
her name was used as an acronym for the purpose of the first project, which was to produce a
(lan)guage (a)nalog of human language in a nonhuman primate
the lana project was established in 1971 by duane rumbaugh, and began with lana
taught a system of communication using keyboard lexigrams (yerkish)
if lana wanted a particular food or rewards, she had to ask computer to give it (ex: please
machine give apple, please machine show slides)

kanzi the bonobo

(sue savage-rumbaugh)
kanzi’s novel sentences: http://www.youtube.com/watch?v=2Dhc2zePJFE
kanzi the toolmaker: http://www.youtube.com/watch?v=1zsSH9UUQtQ

dolphins
(lou herman)
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• http://www.youtube.com/watch?v=jz3sQsTE5tA

alex the african grey parrot

(irene pepperberg)
• http://www.youtube.com/watch?v=ldYkFdu5FJk
• http://www.youtube.com/watch?v=sYk-wE18BTo

kaminski et al., science 2004

during speech acquisition, children form quick and rough hypotheses about the meaning of a
new word after only a single exposure—a process dubbed “fast mapping”
• rico, a border collie, is able to fast map — he knew the labels of over 200 different items
• rico infers the names of novel items by exclusion learning and correctly retrieves those
items right away and 4 weeks later
http://www.sciencemag.org/content/suppl/2004/06/08/304.5677.1682.DC1/1097859s2.mov
• the movie shows the very first session of the identification task, in which a novel item is
requested by using a novel name
• rico is first instructed to bring two familiar items: 1st the “t-rex” (the blue dinosaur), then
“weihnachtsmann” (the little red doll)
• subsequently, a novel word (“sirikid”) is used to ask for the novel item, the white bunny
https://www.youtube.com/watch?v=_6479QAJuz8

animals may be smart, but are they just sort of intellectually-weak humans? are chimps just on
the bottom rung of the ladder up to humans?
“sometimes the human cognitive psychologists can be so fixed on their definitions that they
forget how fabulous these animal discoveries are. we're glimpsing intelligence throughout the
animal kingdom, which is what we should expect. it's a bush, not a single-trunk tree with a line
leading only to us” (clive wynne)
http://www.youtube.com/watch?v=TC1nJ61l-h4
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summary
as humans, we implicitly apply our ‘theory of mind’ to explain animal’s behavior, i.e., we
anthropomorphize
if we want to test the hypothesis that animal thinking is like ours, we should not begin by
assuming that the hypothesis is true!
we don’t always need mental concepts to explain behavior. if a behavior is complex, it does not
necessarily require a mentalistic explanation
if an animal does something in a mechanical way (e.g., copperhead, robin, langur), this doesn’t
mean that they are behaving non-adaptively (e.g., copperhead search image)
thought experiment: how would we do if dropped into animal’s world without the benefits of
cultural support?