Name:______________________________________________

NPB 102, Animal Behavior

Winter 2011, Exam 1

Student ID #_____________________________

Instructions: This exam has 5 pages, none of them blank. Make sure that your copy is complete before
beginning. Write your name on every page and if you want us to be able to return your exam to you by leaving it
in the hall boxes in Briggs hall, sign the authorization at the end of the exam. Read each question very carefully
to be certain that you understand what is being asked.
We will not answer any questions about the exam during the exam! Please do not come up to ask an instructor
about any exam questions during the exam. If you believe a question is a bad question you may make a note on
your exam for the instructors to consider.
Multiple Choice: (1 pt each) Circle the letter of the one best choice to complete each of the following:
1) With respect to song development, the term overproduction refers to:
a) The fact that young male songbirds sing more during the sensory-motor phase than during the sensory
phase.
b) The fact that mature males sing more than young males.
c) The fact that mature males with high testosterone sing more than mature males with low testosterone.
d) The fact that young males in the sensory-motor phase tend to sing more different sounds and/or song
types than they do once they have crystallized their songs.
e) The fact that mature males in the crystallization phase tend to sing more stereotyped songs than they do
during the sensory-motor phase.
2) With respect to song learning, the term sensory phase refers specifically to:
a) The fact that young male songbirds are sensitive to auditory feedback when practicing song during the
sensory-motor phase of song learning.
b) The fact that young male songbirds are sensitive to the acoustic input they receive from singing neighbor
males during territory establishment.
c) The fact that young male songbirds can readily memorize the songs they hear during the first few weeks
of life, but they lose the ability to do this after a certain age.
d) The fact that mature male songbirds can adjust how they respond to neighbor males as a function of what
songs those neighbor males choose to sing.
e) All of the above.
3) Differences in mating system (i.e., monogamy versus polygyny) between prairie and meadow voles could be
because:
a) Males of the two species possess different forms of the receptor for AVP in specific regions of their
brains.
b) Meadow voles evolved under conditions of high density, favoring a polygynous lifestyle, while prairie
voles evolved under conditions of low density, favoring a monogamous lifestyle.
c) Meadow and prairie voles are relatively distantly-related taxa and consequently are unlikely to share
such traits as mating system.
d) All of the above.
e) None of the above.

4) Which of the following procedures would have the most dramatic effect on song if performed on an adult
male songbird such as a zebra finch or white-crowned sparrow?
a) Unilaterally (on one side) lesion (destroy) Area X or lMAN.
b) Bilaterally (on both sides) lesion Area X or lMAN.
c) Bilaterally lesion HVC or RA.
d) Unilaterally lesion nXIIts.
e) Unilaterally lesion HVC or RA.
5) A male songbird that is deafened after tutoring is complete but before it has had the opportunity to practice
its song:
a) Will crystallize an extremely abnormal song that lacks species-typical phonation and syntax.
b) Will not crystallize a song at all.
c) Will crystallize a normal song that is a good match of the tutor songs it heard before it was deafened.
d) Will crystallize a song with species-typical syntax but that is simpler than natural songs.
e) None of the above.
6)

Which of the following would lead you to conclude that inter-individual variation in song in a particular
species was NOT indicative of song dialects?
a) The variants are discretely different from one another, rather than varying on a continuum.
b) The individuals singing different variants are mixed together within localities.
c) The geographic boundaries between groups of birds singing different song variants are abrupt, rather
than gradual.
d) All of the above are consistent with the variation representing dialects.
e) None of the above is consistent with the variation representing dialects.

7) The two voices in the syrinx are:

a) The left and right pairs of medial and lateral labia located at the top of each bronchus.
b) The left and right medial tympanic membranes
c) The two sides of the intraclavicular air sac
d) The left and right syringial musculature
e) The left and right tracheal cartilages
8) The syrinx can produce sound when:
a) The birds air sacs force air outward through one partially-occluded bronchus and the other bronchus is
completely occluded.
b) The birds air sacs force air outward through two fully open bronchi.
c) The birds air sacks forcefully draw air inward through the two sides of the syrinx.
d) The birds air sacs force air outward through one partially-occluded bronchus and the other bronchus is
fully open.
e) None of the above.
9)

Which of the following most accurately describes gene expression patterns in the song system of developing
male songbirds?
a) All of the genes related to song memorization begin to be transcribed throughout the song control nuclei
at about the same time during development.
b) Gene transcription begins at one developmental stage in the motor pathway, and at another
developmental stage in the learning circuit.
c) Gene transcription in the song system is complete by the time young males leave the nest.
d) Gene transcription in the song system does not begin until males begin to crystallize their songs.
e) The developmental stage when gene transcription begins varies among different genes, and among
different song control nuclei.

Name:______________________________________________
Distinguishing Proximate and Ultimate Hypotheses: Each of the following observations and questions is
followed by a list of hypotheses. Indicate what kind of hypothesis each is with a P or U for Proximate or
Ultimate, as well as a D (developmental: How does it Develop?), C (causal: What Causes it?), E (evolutionary
history: How did it Evolve?) or F (fitness: What is its Fitness value?). So, for instance, if you think one of them
is Proximate and Causal, you would write P, C in front of it.

Observation and question 1 (8 pts): Males of a species of blackbird called a common grackle sing only a single
song type despite the fact that female grackles tested under experimental conditions turn out to be more sexually
responsive to multi-song repertoires than to single-song repertoires. Why do male grackles only display
repertoires of a single song-type?
UE

1. The immediate ancestors of common grackles lacked multi-song repertoires, and common grackles
have retained this trait.

PC

2. Male common grackles only have the instructions for a single song type stored in their song control
nuclei.

PD

3. Male common grackles only ever hear a single song type during early life.

UF

4. Rival males are more likely to keep away from females defended by males singing a single song type
than from females defended by males singing multi-song reperoires.

Observation and question 2 (14 pts): Gulls remove the broken eggshells from their nests shortly after the eggs
hatch. Why do they do this?
PC

1. The sight of broken eggshells activates a motor pattern that includes picking up and carrying the
broken shells away from the nest.

PC

2. The smell of the broken shells activates a motor pattern that includes picking up and carrying the
broken shells away from the nest.

UF

3. Proximity of the newly-hatched young to broken eggshells increases the likelihood that they will be
detected and eaten by predators.

UF

4. Proximity of the newly-hatched young to broken eggshells increases the likelihood that they will
become ill through exposure to bacteria that grow on the shells.

UE

5. The immediate ancestor of modern gulls also had evolved to remove eggshells from the nests, and
modern gulls inherited this trait from this ancestor.

PD

6. Newly-hatched gulls observe their parents removing the eggshells and from this experience learn to
remove eggshells themselves.

PD

7. Gulls possess an innate inclination to pick up and carry away broken eggshells.

Paragraph answers: Answer each of the following with a paragraph. Be sure to read the questions carefully,
and keep your answers closely focused on what the question is asking.

1. (5 pts) Describe the evidence that female red-winged blackbirds are more discriminating when it
comes to song than males are, and provide one ultimate explanation for this difference.
You needed to say that female redwinged blackbirds show much more frequent
sexual displays in response to real male redwinged blackbird song than they do to
mockingbirds copies of male redwinged blackbird song, whereas males show
very similar levels of aggressive response (songs and song-spreads) to real male
redwinged blackbird song and mockingbird copies. You could draw a graph
illustrating this if you wanted as part of your answer, but you did not need to do
this in order to get full credit, and verbal description of what the graph means
would be required for full credit.
One likely ultimate explanation for this is that the cost to females of displaying to
the wrong signal could be very high, since it could lead to her copulating with
an inappropriate partner (perhaps even the wrong species) and a dramatic
reduction in her reproductive success. In contrast, the cost to males of NOT
responding to what MIGHT be a genuine competitor would be high, since it could
lead to him losing control of paternity in his mates nest.

2. (6 pts) How do the songs of male great tits differ between populations living in densely-vegetated forests
versus relatively open woodlands? Explain how the effects of these different habitats on signal transmission
could provide the basis for selection leading to the evolution of these differences in song characteristics in
these different habitats.

Woodland great tits sing songs have more high frequency elements in them
than do forest great tits. One hypothesis that could explain this difference is that
the forest and woodland environments differentially affect the propagation of
sounds of different pitch and this leads to selection favoring individuals that
produce the type of song that works best for communication in their particular
habitat. Specifically, high pitched sounds do not propagate as well in physically
complicated habitats such as forest as they do in simpler, more open habitats such
as woodlands. Consequently, individuals singing songs emphasizing high
frequencies would be selectively disfavored in forest compared with individuals
whose songs contained primarily low-frequency notes that propagate better there.
You could get up to 5 points if your answer goes only this far. For the last point
you would need to at least make an attempt to explain why high frequency notes
should be so widespread in songs of woodland great tits. It would be sufficient
just to say that the lack of selection against use of high frequency notes in
woodlands would leave individuals in these populations free to use high
frequency notes.

Name:______________________________________________
3. (8 pts) Some species of songbirds that display multi-song repertoires, such as song sparrows,
can escalate or defuse aggressive encounters through the songs they select when responding
to a singing neighbor. Explain what it means for a song sparrow to respond to a singing
neighbor with (1) a song type match, (2) a song repertoire match, or (3) an unshared song,
and then explain how responding with a song type match, a repertoire match, or an unshared
song would influence whether the aggressive interaction then escalates (becomes more
aggressive), remains about the same, or is defused (becomes less aggressive).
First you needed to explain the three kinds of responses. (1) A song type match is the
term applied when the focal male responds with a song that is an extremely close match
to the song sung by his opponent. That is, he responds with a song that is the same type
as the one sung at him. (2) A song repertoire match is the term applied when the focal
male response with a song that is of a different type from the one his opponent sang at
him, but it is of a type that the opponent does have in his repertoire. That is, it matches
something in his opponents repertoire, but it is not of the same type as the song the
opponent originally sang. (3) An unshared song is just a song type that the opponent
doesnt even have in his repertoire.
Next you needed to explain how these different responses could be used to escalate or
defuse an aggressive interaction.
If a male responds to an opponents song by singing a type match, this is the most
aggressive response and is the most likely response to cause escalation to physical
violence.
If a male responds to an opponents song by singing a repertoire match, this is an
intermediate type of response that does not specifically escalate or defuse the interaction.
(That would be enough to say about this, but if you wanted you could go on to note that
whether this leads to escalation or not depends on how the original male now responds
with a type match, or with another repertoire match, or with an unshared song.)
If a male responds to an opponents song by singing an unshared song, this is a
minimally aggressive response, and it is most likely to defuse or end the interaction
without physical violence.

I hereby give the instructors of NPB 102 my permission to return my exam to me by leaving it in the hall boxes
in Briggs Hall.
Signature: ______________________________________________________________