NPB 102, Animal Behavior

Fall 2012
HAHN Study Questions for last section of course
12. What is a search image? PERCEPTUAL SCREENING MECHANISM THAT
IMPROVES DETECTION OF ITEMS THAT POSSESS PARTICULAR
CHARACTERISTICS, USUALLY FOODS. What advantages (benefits) do
search images confer IMPROVES PREY DETECTION EFFICIENCY, and what
potential disadvantages (costs) CAN CAUSE SOME DESIRABLE PREY THAT
DONT MATCH SEARCH IMAGE TO BE OVERLOOKED/MISSED?
Describe experimental evidence illustrating these advantages and disadvantages.
EXAMPLE FROM LECTURE IS ON PORTIA SPIDERS, WHICH ARE MORE
LIKELY TO DETECT THE PREY SPECIES THEY ARE PRE-EXPOSED TO
THAN THEY ARE ANOTHER EQUALLY-SUITABLE PREY SPECIES THAT
THEY WERE NOT PRE-EXPOSED TO. SO WHEN THEY HAVE A
SEARCH IMAGE THEIR SEARCHING EFFICIENCY IS HIGH (BENEFIT)
FOR THE PREY THAT MATCHES THE IMAGE, BUT RELATIVELY LOW
(COST) FOR THE OTHER. THEY DONT DEVELOP A SEARCH IMAGE
FOR PRE-EXPOSED PREY IF IT WAS A TYPE THAT THEY DONT
NORMALLY PREFER (E.G., FLIES), SO THIS ONLY APPLIES TO PREY
THEY ACTUALLY NORMALLY PREFER TO EAT.
13. What happens when Portia spiders are first primed with exposure to a preferred
prey item and then presented with either the same or different prey in (a) a test
situation where the prey is freely visible THEY ARE EQUALLY LIKELY TO
DETECT ANY PREY, EVEN A NON-PREFERRED PREY LIKE A FLY, IF
THEY WERE PRIMED BY EXPOSURE TO A PREFERRED PREY ITEM
AND THEN PRESENTED WITH FREELY-VISIBLE PREY, and (b) a test
situation where the prey is partially concealed THEY ARE BETTER AT
DETECTING THE PREY TYPE THEY WERE PRE-EXPOSED TO THAN
THE TYPES THEY WERE NOT PRE-EXPOSED TO IF THE PRE-EXPOSURE
PREY IS A PREFERRED PREY TYPE AND THE TEST PREY ARE
PARTIALLY CONCEALED? What does this result reveal about the role of
search images in foraging behavior ITS CONSISTENT WITH THE
INTERPRETATION THAT SEARCH IMAGES FUNCTION MAINLY TO
HELP THE ANIMAL LOCATE PREY UNDER CHALLENGING
CONDITIONS, SUCH AS WHEN THE PREY ARE HIDDEN, NOT WHEN
DETECTING THE PREY IS EASY. NOTE ALSO THAT THE FACT THAT
EVEN NON-PREFERRED PREY ARE READILY FIXED ON (DETECTED)
WHEN THE PRIMING PREY WAS A PREFERRED PREY TYPE AND
WHEN THE TEST PREY PRESENTED IS FREELY VISIBLE SUGGESTS
THAT EXPOSURE TO THE PRIMING PREY ALSO HAS A GENERAL
EFFECT ON MOTIVATION TO SEARCH, NOT ONLY ON THE
ATTRIBUTES OF THE SEARCH IMAGE.?
14. Explain whether you think it is more useful to quantify detection or
consumption when evaluating whether or not a search image is present. THE
SEARCH IMAGE IS USED IN PREY DETECTION, NOT IN THE DECISION

WHETHER TO EAT THE PREY ONCE IT IS DETECTED. ONCE THE

FORAGER LOCATES A PROSPECTIVE PREY ITEM USING A SEARCH
IMAGE, IT THEN MAY EVALUATE THE PREY MORE CAREFULLY
USING MORE DETAILED CRITERIA AND DECIDE ON THAT BASIS
WHETHER TO ACTUALLY EAT IT OR NOT. SO DETECTION IS A
BETTER THING TO QUANTIFY THAN CONSUMPTION WHEN
EVALUATING WHETHER A SEARCH IMAGE IS PRESENT.
15. What is frequency-dependent selection, and how can it account for the coexistence of alternate foraging phenotypes? FREQUENCY-DEPENDENT
SELECTION IS SELECTION UNDER WHICH THE FITNESS OF A
PARTICULAR TRAIT DEPENDS ON THE PREVALENCE OF ANOTHER
TRAIT IN THE POPULATION. Provide at least one specific example. THE
PERISODUS CICHLID FISH ARE WELL-DESCRIBED IN THE BOOK.
THEY HAVE ASYMMETRIC JAWS THAT FACILITATE ATTACKING
OTHER FISH FROM THE SIDE (THEY EAT SCALES OFF THE FLANKS OF
OTHER FISH). WHEN THE PROPORTION OF EACH MORPH (LEFT- AND
RIGHT-JAWED INDIVIDUALS) IN THE POPULATION IS ABOUT EQUAL,
THEN THE FITNESSES OF THESE TWO MORPHS ARE ALSO EQUAL.
THIS IS PROBABLY BECAUSE (A) THE SIDE THAT THE FISH HAS TO
ATTACK FROM IS DETERMINED BY THE JAW MORPHOLOGY (A
RIGHT-JAWED PERISODUS MUST ATTACK FROM THE LEFTTHEY
ALWAYS APPROACH FROM BEHIND THE VICTIM), AND (B) THE PREY
FISH WILL BE MORE VIGILANT ABOUT ATTACKS COMING FROM THE
SIDE THAT THEY ARE MOST FREQUENTLY ATTACKED FROM, SO
WHICHEVER JAW MORPH IS MORE COMMON IN THE POPULATION
WILL BE AT A DISADVANTAGE COMPARED WITH THE OTHER.
SOIF THE PROPORTION OF LEFT- AND RIGHT-JAWED PERISODUS
HAPPENS TO DRIFT AWAY FROM 50:50, WHICHEVER IS LESS
COMMON WILL HAVE HIGHER FITNESS IN THAT GENERATION AND
THE NUMBERS OF THAT MORPH WILL THEREFORE INCREASE IN THE
NEXT GENERATION, SHIFTING THE PROPORTION BACK TOWARD
50:50. ACTUAL DATA COLLECTED ON PERISODUS (SHOWN IN THE
BOOK) SUPPORT THIS.
ANOTHER EXAMPLE THAT IS IN THE LECTURE SLIDES BUT NOT IN
THE BOOK IS ABOUT ROSEATE TERNS. TERNS NORMALLY CATCH
FISH BY DIVING INTO THE WATER AND GRABBING THEM IN THEIR
BILLS. SOME POPULATIONS OF ROSEATE TERNS HAVE A SMALL
PROPORTION OF INDIVIDUALS THAT NEVER HUNT BUT INSTEAD
ALWAYS JUST STEAL FISH THAT OTHER INDIVIDUALS HAVE
CAUGHT. THEY STEAL THEM IN FLIGHT. THAT IS, THEY SEE
ANOTHER ROSEATE TERN THAT HAS A FISH IN ITS BILL AND THEY
CHASE IT AND ATTEMPT TO TAKE THE FISH AWAY. THESE TWO
FEEDING TACTICS PERSIST IN THE POPULATION, BUT THE NUMBER
OF INDIVIDUALS THAT USE THE STEAL FISH TACTIC NEVER GETS
VERY HIGH. THE QUESTION IS WHY DO THE TWO TACTICS BOTH
PERSIST IN THE POPULATION (RATHER THAN ONE TACTIC

OUTCOMPETING THE OTHER AND GOING TO FIXATION WHILE THE

INFERIOR TACTIC GOES TO EXTINCTION), AND GIVEN THAT THEY
DO PERSIST, WHY AT THE PARTICULAR RELATIVE FREQUENCIES
THAT THEY DO? FREQUENCY-DEPENDENT SELECTION IS THE
ANSWER. IN THE FIGURE INCLUDED IN THE LECTURE NOTES, THE Y
AXIS SHOULD BE RELATIVE, NOT ABSOLUTE, FITNESS OF EACH
PHENOTYPE. THE WAY TO READ IT IS TO IMAGINE A POPULATION
THAT INITIALLY HAS NO STEALERS I.E., A POPULATION WHERE
ALL INDIVIDUALS OBTAIN FOOD BY HUNTING/CATCHING FISH. IF A
FEW INDIVIDUALS THAT INSTEAD STEAL FISH APPEAR IN THE
POPULATION, THEIR PROPORTION IN THE POPULATION WILL
INITIALLY BE VERY LOW. THEY WILL THEREFORE HAVE ACCESS TO
AN ALMOST UNLIMITED SUPPLY OF THE HUNTERS WHO ARE
DOING THE WORK OF CATCHING THE FISH, AND FURTHER, THOSE
HUNTERS WILL SELDOM ENCOUNTER THE STEALERS (BECAUSE
THERE ARE, PROPORTIONALLY, VERY FEW STEALERS), SO THEY ARE
NOT LIKELY TO BE VERY VIGILANT ABOUT PROTECTING THE FISH
THEY HAVE JUST CAUGHT FROM BEING STOLEN (I.E., THE STEALER
WOULD ZIP IN AND TAKE IT AWAY BEFORE THE HUNTER EVEN
KNOWS WHATS HAPPENING). UNDER THESE CIRCUMSTANCES, THE
FITNESS OF THE STEALING PHENOTYPE IS LIKELY TO BE VERY
HIGH RELATIVE TO THAT OF THE HUNTING PHENOTYPE, BECAUSE
THE STEALERS ARE GETTING FOOD MORE QUICKLY AND EASILY
THAN THE HUNTERS ARE. SINCE THIS IS AN INHERITED PHENOTYPE,
OVER THE COMING GENERATIONS THE PROPORTION OF STEALERS
IN THE POPULATION WILL INCREASE (MOVING TO THE RIGHT FROM
THE ORIGIN IN THE FIGURE IN THE LECTURE NOTES). HOWEVER, AS
THE PROPORTION OF STEALERS IN THE POPULATION CONTINUES TO
INCREASE, THIS WILL NECESSARILY LEAD TO AN INCREASE IN
COMPETITION AMONG THE STEALERS FOR THE FISH THAT HAVE
BEEN CAUGHT BY HUNTERS, AS WELL AS THE EDUCATION OF THE
HUNTERS SO THAT THEY BECOME HARDER TO STEAL FROM (I.E., IF
A HUNTER HAS RECENTLY HAD A FISH STOLEN, IT IS VERY LIKELY
TO BE MUCH MORE VIGILANT ABOUT WATCHING OUT FOR
STEALERS AND BETTER AT PREVENTING ITS HARD-WON PREY FROM
BEING STOLEN IN FUTURE). CONSEQUENTLY, AT SOME POINT THE
PROPORTION OF STEALERS IN THE POPULATION WILL GET HIGH
ENOUGH THAT IT IS NO EASIER TO GET PREY BY STEALING THAN
IT IS TO GET IT BY HUNTING. THE PROPORTION OF STEALERS AND
HUNTERS IN THE POPULATION WILL THEN HOVER AROUND THIS
EQUILIBRIUM VALUE BECAUSE ANY TIME IT DRIFTS TO A HIGHER
PROPORTION OF ONE THAN THE OTHER, THE PHENOTYPE THAT IS
RARER THAN THE PROPORTION DICTATED BY THIS EQULIBRIUM
WILL HAVE HIGHER FITNESS THAN THE OTHER AND WILL
THEREFORE INCREASE IN THE NEXT GENERATION, THEREBY
BRINGING THE POPULATION BACK TOWARD THIS EQULIBRIUM.

NOTE THAT UNLIKE IN THE PERISODUS EXAMPLE, THE

PROPORTIONS OF THE TWO PHENOTYPES IN THE POPULATION AT
WHICH EQUAL FITNESSES OCCUR IS NOT EXPECTED TO BE 50:50,
BUT AT SOME QUITE LOW PREVALENCE OF STEALERS. THE EXACT
PROPORTION WHERE THE EQUILIBRIUM WILL BE REACHED WILL
DEPEND ON A NUMBER OF THINGS, SUCH AS THE ABUNDANCE OF
FISH (I.E., HOW EASY IS IT TO SUCCEED BY HUNTING?), AND HOW
EASY OR DIFFICULT IT IS FOR THE HUNTERS TO PROTECT
THEMSELVES FROM HAVING FISH STOLEN ONCE THEY LEARN THAT
STEALERS DO EXIST IN THE POPULATION, BUT IN GENERAL THE
BALANCE POINT WILL BE REACHED AT A RELATIVELY LOW
PROPORTION OF STEALERS IN THE POPULATION.
REMEMBER
THAT IN GENERAL, FREQUENCY-DEPENDENT SELECTION IS
INVOKED AS A POSSIBLE EXPLANATION FOR TWO OR MORE
PHENOTYPES CO-EXISTING IN A POPULATION. IF THE SELECTION
WAS ABSOLUTE RATHER THAN FREQUENCY-DEPENDENT,
WHICHEVER PHENOTYPE WAS SUPERIOR WOULD SIMPLY EXCLUDE
THE OTHER(S) FROM THE POPULATION, AND ONLY ONE TACTIC
WOULD PERSIST OVER THE LONG TERM.
16. What is the difference between a strategy and a tactic? STRATEGIES ARE
THE SET OF RULES THAT INDIVIDUALS FOLLOW TO DETERMINE
WHAT BEHAVIORAL TACTICS THEY WILL USE. TACTICS ARE THE
ACTUAL BEHAVIORS THEY ENGAGE IN. Give an example where (a) a
single tactic defines a strategy, IN FIXED STRATEGIES AN INDIVIDUAL
WILL ENGAGE IN ONLY A SINGLE TACTIC (E.G., BE A TERRITORIAL
MALE, OR BE A FISH STEALER), AND THE RULE IT FOLLOWS WILL BE
UNCONDITIONAL: I WILL ALWAYS TRY TO BE TERRITORIAL AND
NEVER BE A SATELLITE MALE OR I WILL ALWAYS ATTEMPT TO
STEAL FISH AND NEVER HUNT FOR THEM MYSELF. THE ROSEATE
TERNS PROVIDE ONE EXAMPLE: INDIVIDUALS FOLLOW ONLY ONE
RULE (EITHER I WILL HUNT OR I WILL STEAL) AND THEREFORE
ENGAGE IN ONLY ONE BEHAVIORAL TACTIC (HUNT FOR FISH OR
STEAL FISH) THROUGHOUT THEIR LIVES. THE PERISODUS FISH
PROVIDE ANOTHER EXAMPLE (THE RULES BEING I WILL ALWAYS
ATTACK FROM THE LEFT, OR I WILL ALWAYS ATTACK FROM THE
RIGHT, WITH THE TACTICS BEING ATTACK FROM LEFT OR
ATTACK FROM RIGHT) and (b) a strategy involves multiple tactics.
CONDITIONAL STRATEGIES INVOLVE RULES THAT SAY THINGS LIKE
IF SUCH AND SUCH IS TRUE THEN I WILL DO THIS, ELSE I WILL DO
THAT. UNDER THESE RULES, THE INDIVIDUAL HAS A CHOICE
BETWEEN TWO OR MORE TACTICS, AND THE CIRCUMSTANCES WILL
DETERMINE WHICH TACTIC IS USED. SO HERE, THE STRATEGY IS
NOT DEFINED BY A SINGLE TACTIC, BUT BY A COMBINATION OF
POSSIBLE TACTICS. EXAMPLES INCLUDE THE RUDDY TURNSTONES,
WHICH FOLLOW THE RULES (STRATEGY): I WILL DEFEND AND FEED
FROM A KELP PILE IF I CAN SUCCESSFULLY ACQUIRE ONE, ELSE I

WILL NOT BE TERRITORIAL AND JUST FEED OUT ON THE OPEN

BEACH SAND. UNLIKE IN FIXED STRATEGIES (LIKE THE ROSEATE
TERN AND THE PERISODUS), YOU SHOULD EXPECT INDIVIDUALS
EMPLOYING CONDITIONAL STRATEGIES TO BE ABLE TO SWITCH
BETWEEN TACTICS, AND THAT THE FITNESSES OF THE
INDIVIDUAL TACTICS SHOULD BE DIFFERENT. THAT IS,
FORAGING EFFICIENCY OF TURNSTONES USING THE DEFEND A
KELP PILE TACTIC WILL BE HIGHER THAN FOR THOSE USING THE
OPEN BEACH FORAGING TACTIC. THE ONLY REASON THE ONES USE
THE INFERIOR TACTIC IS THAT THEYRE MAKING THE BEST OF A
BAD SITUATION. IF THEY TRIED TO DEFEND A KELP PILE INSTEAD,
THEY WOULD DO EVEN WORSE THAN IF THEY HUNT FOR LOOSE
THINGS OUT ON THE OPEN BEACH SAND BECAUSE THEY WOULD
FAIL TO GET A KELP PILE (THEY MIGHT EVEN HAVE ZERO
FORAGING SUCCESS BECAUSE THEY SPEND ALL THEIR TIME
TRYING AND FAILING TO GET A KELP PILE INSTEAD OF FORAGING).
17. Describe the behavior of ruddy turnstones foraging on sandy beaches with
specific reference to the different tactics that individuals use. Is each of these
tactics what you would call a distinct strategy or just a separate tactic of a
single conditional strategy? Explain your reasoning. I ANSWERED THIS
ABOVE.
18. What would you need to know in order to determine whether fish stealing
behavior by roseate terns is a tactic of a conditional strategy or a fixed strategy?
IT WOULD HELP TO KNOW ALL OF THE FOLLOWING: CAN
INDIVIDUALS SWITCH FROM ONE TACTIC TO THE OTHER? IS THE
FORAGING SUCCESS (A PROXY FOR FITNESS) THE SAME OR
DIFFERENT FOR THE TWO TACTICS? DOES THE PREVALENCE OF THE
TWO TACTICS REMAIN SIMILAR ACROSS A NUMBER OF
GENERATIONS? SEE ANSWER TO #5 ABOVE FOR MORE INFO ON
THIS.
19. Explain how optimality models have been used to better-understand how crows
foraging on whelks choose and handle prey. THE EXAMPLE IN THE BOOK
EXPLAINS CLEARLY HOW THE ISSUES OF WHELK SELECTION, HOW
HIGH CROWS FLY TO DROP THEM, AND HOW MANY TIMES THEY
DROP THEM BEFORE GIVING UP FIT WITH OPTIMAL FORAGING
THEORY. THE MODELS PREDICT THAT THE CROWS SHOULD SELECT
THE SIZES OF WHELKS THAT GIVE THEM THE GREATEST
ENERGY/NUTRITION RETURN PER UNIT EFFORT. IN THIS CASE THE
CROWS SELECT ONLY THE LARGEST WHELKS, AND CARRY THEM
ABOUT 5 METERS UP INTO THE AIR, ON AVERAGE, BEFORE
DROPPING THEM. THEY ALSO KEEP DROPPING THE SAME WHELK
UNTIL IT BREAKSTHEY DONT GIVE UP AND SWITCH TO A
DIFFERENT WHELK IF ONE DOESNT BREAK AFTER A FEW DROPS.
EXPERIMENTAL TESTS OF HOW LONG IT TAKES TO BREAK WHELKS
OF DIFFERENT SIZES FROM DIFFERENT HEIGHTS SHOW THAT LARGE
WHELKS BREAK MORE EASILY THAN SMALLER ONES OVERALL

(PROBABLY BECAUSE THEY HIT THE GROUND WITH MORE FORCE

BECAUSE THEY WEIGH MORESEE THE FIGURE IN TEXT AND
LECTURE SLIDES), THE LIKELIHOOD THAT ONE WILL BREAK ON ANY
GIVEN DROP IS INDEPENDENT OF HOW MANY TIMES IT HAS
ALREADY BEEN DROPPED (SO SWITCHING TO ANOTHER WHELK IS
UNLIKELY TO BE A GOOD IDEA), AND CARRYING THEM HIGHER
THAN ABOUT 5 METERS DOES NOT IMPROVE THE CHANCES OF
THEM BREAKING (SEE THE FIGURE IN THE TEXT AND LECTURE
SLIDES). How have those models been fine-tuned using empirical data? I
MOSTLY ANSWERED THIS ABOVE (EMPIRICAL DATA ON WHAT IT
TAKES TO BREAK A WHELK, FOR INSTANCE). ADDITIONAL
ESTIMATES HAVE BEEN MADE OF HOW MUCH ENERGY THE CROWS
WOULD GET IF THEY FORAGED ON SMALL, MEDIUM AND LARGE
WHELKS AND WHEN THE ENERGY AND TIME REQUIRED TO GET THE
FOOD OUT OF THE DIFFERENT SIZE WHELKS IS COMBINED WITH THE
ESTIMATE OF THE ENERGY OBTAINED BY EATING THEM, IT
REVEALS THAT LARGE WHELKS ARE INDEED THE BEST, MEDIUM
ONES WORSE, AND IT ACTUALLY COSTS MORE TO FEED ON SMALL
WHELKS THAN THE CROW GETS BACK FROM EATING THEM.
20. What are the round dance and the waggle dance performed by honeybees,
and how are they thought to facilitate efficient foraging? THESE ARE
DESCRIBED IN DETAIL IN THE TEXTBOOK. ROUND DANCE IS THE
SIMPLE LITTLE RUNNING-AROUND-IN-A-SMALL-CIRCLE DANCE
THAT SOME BEES DO WHEN THEY COME BACK TO THE HIVE. THIS
ONE SIGNALS TO OTHERS THAT THERE IS A RICH NECTAR SOURCE
SOMEWHERE NEAR THE HIVE (E.G., WITHIN, SAY, 50M). IT PROVIDES
NO INFORMATION ABOUT DIRECTION TO THE FOOD SOURCE. THE
WAGGLE DANCE IS MORE COMPLEX AND INCLUDES A STRAIGHT
RUN SECTION IN THE MIDDLE OF SOME LEFT AND RIGHT LOOPS.
DURING THE STRAIGHT RUN PART, THE BEE WAGGLES ITS
ABDOMEN. THIS DANCE PROVIDES ADDITIONAL QUITE COMPLEX
INFORMATION TO THE OTHER BEES OBSERVING IT. THE DIRECTION
OF THE STRAIGHT RUN INDICATES COMPASS DIRECTION TO THE
RICH FOOD SOURCE. IF THE BEE IS DANCING ON A HORIZONTAL
SURFACE, THEN THE DIRECTION IT GOES DURING THE RUN IS THE
COMPASS DIRECTION THE BEES SHOULD FLY WHEN THEY GO
OUTSIDE. IF THE BEE IS DANCING INSIDE THE HIVE IN THE DARK ON
A VERTICAL SURFACE, THEN THE ANGLE THAT THE BEE TAKES
AWAY FROM VERTICAL TELLS THE OTHER BEES WHAT ANGLE
RELATIVE TO THE SUNS POSITION THEY SHOULD FLY WHEN THEY
GO OUTSIDE. STRAIGHT AT THE SUN WOULD BE SIGNIFIED BY A
RUN STRAIGHT UPWARD. DIRECTLY AWAY FROM THE SUN WOULD
BE A RUN STRAIGHT DOWN. ETC. THERE ALSO IS INFORMATION IN
THE LEGNTH OF THE RUN/NUMBER OF WAGGLES, ABOUT THE
DISTANCE TO THE SOURCE. THESE DANCES ARE THOUGHT TO
IMPROVE HIVE PERFORMANCE BY IMPROVING THE ABILITY OF

FORAGERS TO QUICKLY LOCATE AND EXPLOIT NEWLY-EMERGENT

NECTAR/POLLEN SOURCES, SINCE THE INFORMATION SPREADS
QUICKLY FROM THE FORAGERS WHO FIRST DISCOVER THEM TO THE
OTHER FORAGERS IN THE HIVE. Describe an experimental manipulation the
results of which are consistent with this idea. LOTS OF EXPERIMENTS
TESTING THIS HAVE BEEN DONE BUT PARTICULARLY COMPELLING
ONES ARE DESCRIBED IN FIGURES 7.21 AND 7.23 ON PAGES 240-241
OF ALCOCK. THE EXPERIMENT FOR FIGURE 7.23 IS ESPECIALLY
CLEVER. ITS DESCRIBED IN DETAIL IN THE TEXT ON PAGE 241.
VERY BRIEFLY, WORKER BEES WERE FOOLED INTO THINKING
THEY HAD FLOWN A LONG WAY TO THE FOOD SOURCE BY MAKING
THEM FLY DOWN A TUNNEL THAT WAS ACTUALLY SHORT, BUT
THAT HAD A LOT OF VISUAL INFORMATION GOING BY ON THE
WALLS, MAKING THE BEES THINK THEY HAD FLOWN A LONG
WAY. THIS WOULD BE KIND OF LIKE HAVING YOU LOOK OUT THE
SIDE WINDOW OF A CAR YOURE RIDING IN AND TELLING YOU TO
GUESS HOW FAR YOU HAD GONE BASED ON HOW MANY FENCE
POSTS OR TELEPHONE POLES WENT BY IN A PARTICULAR PERIOD OF
TIME, BUT THE CAR IS ACTUALLY GOING SLOWLY AND THE POSTS
ARE BEING MADE TO LOOK LIKE THEYRE GOING BY QUICKLY, SO
YOU THINK YOURE GOING FAST WHEN YOURE ACTUALLY JUST
CRAWLING ALONG. THIS IS ESSENTIALLY WHAT THEY DID TO THE
BEES, SO THEY FLEW A FEW METERS BUT THE VISUAL INFO THAT
WENT BY WAS CONSISTENT WITH THEM HAVING FLOWN A LONG
WAY. WHEN THEY CAME BACK AND DANCED AT THEIR HIVES,
THEY INDICATED WITH THEIR DANCES THAT THE FOOD SHOULD BE
FOUND FAR FROM THE HIVE, AND MOST IMPORTANTLY, THE
WORKERS THAT THEN WENT OUT LOOKING FOR IT WENT TO THE
DISTANCE THAT THE DANCERS HAD INDICATED (70 OR SO METERS
FROM THE HIVE), NOT TO WHERE THE FOOD HAD ACTUALLY BEEN
(ONLY ABOUT 8 METERS AWAY). NOTE THAT THE RECRUITS (THE
BEES THAT OBSERVED THE DANCE) WERE NOT OBLIGED TO FLY
DOWN THE TUNNEL. THEY JUST FLEW OUTSIDE.
21. When we study the ultimate causes of variation in behavior we need to assess
fitness in some way. Ideally, we should assess fitness directly. What do we need
to measure if we want to assess fitness directly? IDEALLY, LIFETIME
REPRODUCTIVE SUCCESS, WHICH IS A COMBINATION OF
INDIVIDUAL REPRODUCTIVE SUCCESSES MULTIPLIED ACROSS THE
ENTIRE LIFETIME OF THE ANIMAL. SOBOTH SURVIVAL AND
REPRODUCTIVE SUCCESS ARE CONSIDERED TO BE DIRECT
MEASURES OF FITNESS. In practice, it is not always practical to measure
fitness directly, in which case we may instead use fitness proxies. What is a
fitness proxy? A FITNESS PROXY IS ANYTHING WE MEASURE THAT
WE BELIEVE IS WELL-CORRELATED WITH FITNESS, EVEN THOUGH IT
IS NOT A DIRECT MEASURE OF FITNESS ITSELF. What are some specific
examples of fitness proxies? (Hint: Think about assumptions underlying

optimal foraging theory.) THERE ARE LOTS. NET ENERGY INTAKE PER
UNIT TIME IS USED A LOT, SINCE IT IS EXPECTED TO BE RELATED TO
FITNESS (INDEED OPTIMAL FORAGING THEORY ASSUMES THAT NET
ENERGY INTAKE PER UNIT TIME WILL IN FACT BE TIGHTLY
CORRELATED WITH FITNESS). OTHERS THAT WE HAVE
ENCOUNTERED DURING THE QUARTER INCLUDE THE NUMBER OF
COPULATION SOLICITATIONS THAT A MALE BIRDS SONG INDUCES
FEMALES TO GIVE (THIS ONE IS FAIRLY CLOSE TO DIRECT MEAUSRE
OF REPRODUCTIVE SUCCESS, BUT REMEMBER THAT A MALE MIGHT
GET LOTS OF COPULATIONS AND YET, FOR SOME REASON SUCH AS
SPERM COMPETITION, FAIL TO ACTUALLY FERTILIZE MANY EGGS),
NUMBER OF MATES A MALE GETS (E.G., WITH THE GREAT REED
WARBLERAGAIN, IF HE FAILS TO SIRE MANY OF THE OFFSPRING
FOR SOME REASON, THE FACT THAT HE HAS MANY MATES MAY
NOT TRANSLATE INTO HIGH REPRODUCTIVE SUCCESS). THERE ARE
MANY OTHERS. What are some advantages and disadvantages of using fitness
proxies instead of directly measuring fitness? ADVANTAGES MAINLY HAVE
TO DO WITH THEM BEING EASIER TO MEASURE THAN FITNESS
ITSELF IS. LIFETIME REPRODUCTIVE SUCCESS IS REALLY HARD TO
MEASURE IN MOST CASES, BUT FITNESS PROXIES LIKE NUMBER OF
MATES ONE GETS ARE LOTS EASIER. Can you think of any advantages to
measuring fitness directly and also measuring some fitness proxies at the same
time? THE MOST OBVIOUS ADVANTAGE IS THAT YOU WILL HAVE
CALIBRATED YOUR FITNESS PROXY SO THAT YOU KNOW,
EMPIRICALLY, WHAT THE RELATIONSHIP BETWEEN YOUR PROXY
AND LIFETIME REPRODUCTIVE SUCCESS IS. BUT THERE IS ALSO THE
POSSIBILITY THAT YOU COULD DETERMINE THE EXTENT TO WHICH
CERTAIN ELEMENTS OF BEHAVIOR, PHYSIOLOGY, WHATEVER
ACTUALLY CONTRIBUTE TO FITNESS. SO, FOR INSTANCE, IF YOU
HAVE A FORAGING SUCCESS FITNESS PROXY AND IT HAS AN
EXTREMELY STRONG CORRELATION WITH ACTUAL FITNESS, YOU
MAY BE ABLE TO MAKE SOME INFERENCES ABOUT HOW
DEPENDENT (AS IN CAUSE-EFFECT) FITNESS IS ON FORAGING
SUCCESS IN YOUR SYSTEM. IF YOU ALSO HAD MEASURED OTHER
FITNESS PROXIES SIMULTANEOUSLY, YOU MIGHT BE ABLE TO
EVALUATE ALTERNATIVE HYPOTHESES ABOUT WHICH SPECIFIC
ACTIVITIES CONTRIBUTE MORE TO FITNESS THAN OTHERS.
22. What does it mean to say that behavior of individuals can affect patterns at
higher levels of organization? WHAT INDIVIDUALS ARE DOING
INFLUENCES PATTERNS WE SEE AT THE POPULATION LEVEL,
COMMUNITY LEVEL, OR EVEN HIGHER LEVELS OF ORGANIZATION.
THESE PATTERNS INCLUDE THINGS LIKE HOW INDIVIDUALS ARE
DISTRIBUTED IN SPACE (ARE THEY CLUSTERED IN GROUPS, OR
EVENLY SPREAD OUT ON SEPARATE TERRITORIES, OR DO THEY
SPEND PART OF THE YEAR IN ONE KIND OF GROUPING AND PART
ANOTHER? ETC), AND HOW PARTICULAR SPECIES ARE DISTRIBUTED

IN SPACE AND TIME (I.E., COMMUNITY-LEVEL PATTERNS). How does

work on oystercatchers and high-intertidal limpets illustrate this idea? THERE
ARE SEVERAL ELEMENTS TO THIS. SELECTIVE OYSTERCATCHER
PREDATION CAN HAVE A DIRECT EFFECT ON WHERE LIMPETS OF
PARTICULAR SPECIES AND SIZES ARE FOUND, AND ALSO CAN
IMPOSE SELECTION ON LIMPET BEHAVIOR THAT FURTHER
CONTRIBUTES TO THESE PATTERNS. THE OYSTERCATCHER
PREFERENCE FOR DIGITALIS OVER SCABRA COMBINED WITH THE
OYSTERCATCHER NEGLECT OF LIMPETS SMALLER THAN 10MM
LENGTH LEADS TO DIGITALIS LARGER THAN 10MM BEING
RESTRICTED TO VERTICAL SURFACES OUT OF REACH OF
OYSTERCATCHERS, AND LEAVES SCABRA ESSENTIALLY FREE TO
OCCUPY ALL OTHER SURFACES THAT ARE OTHERWISE PHYSICALLY
ACCEPTABLE TO THEM (IN TERMS OF EXPOSURE TO SEA WATER
AND AIR AND HEAT AND COLD AND WHATNOT). THE BEHAVIOR OF
THE LIMPETS CONTRIBUTES BECAUSE ITS SCABRAS HOMING
BEHAVIOR THAT MAKES THEM SO CHALLENGING FOR
OYSTERCATCHERS TO REMOVE, THEREBY MAKING
OYSTERCATCHERS NEGLECT THEM. AND DIGITALIS UPWARDMIGRATORY BEHAVIOR CONTRIBUTES TO THE DISTRIBUTION
PATTERN BY INCREASING THE LIKELIHOOD THAT LARGE DIGITALIS
WILL BE FOUND HIGH ON THE SHORE ON VERTICAL SURFACES (I.E.,
REINFORCES THE DIRECT EFFECT THAT OYSTERCATCHER
PREDATION IS HAVING ON THIS).
23. What is intertidal zonation and what kinds of factors are thought to be
responsible for it? ORGANISMS OF PARTICULAR TYPES ARE
CLUSTERED IN ZONES RELATIVE TO TIDAL HEIGHT. PHYSICAL
FACTORS SUCH AS AMOUNT OF EXPOSURE TO AIR (MORE THE
HIGHER YOU GO ON THE SHORE) OR SEA WATER (MORE THE LOWER
YOU GO), HEAT AND COLD, AND WAVE ACTION ALL ARE VERY
IMPORTANT. BIOTIC FACTORS SUCH AS PREDATION AND
COMPETITION FOR SPACE OR FOOD ALSO CONTRIBUTE.
24. What is the difference between absolute and relative intertidal height?
ABSOLUTE HEIGHT IS THE ABSOLUTE POSITION, IN METERS OR FEET
OR WHATEVER, RELATIVE TO THE MEAN LOWER LOW WATER
LEVEL MEASURED EMPIRICALLY FOR A PARTICULAR PLACE ON A
COAST. IT IS WHERE THE WATER WOULD BE IF THE TIDE WERE AT
THAT HEIGHT AND THERE WAS ABSOLUTELY NO WAVE ACTION AT
ALL. RELATIVE TIDAL HEIGHT TAKES INTO ACCOUNT VARIATION
IN THE PHYSICAL FACTORS THAT OCCURS BECAUSE OF
DIFFERENCES AMONG SITES IN WAVE ACTION, WEATHER, AND
WHATNOT. SO FOR INSTANCE, TWO LOCATIONS NEAR EACH OTHER
THAT ARE AT EXACTLY THE SAME ABSOLUTE TIDAL HEIGHT (LETS
SAY +1 METER) COULD BE AT VERY DIFFERENT RELATIVE TIDAL
HEIGHTS. IF ONE OF THE SITES IS, SAY, EXPOSED TO HEAVY WAVE
ACTION, THEN THE MARINE ENVIRONMENT REACHES UP TO A

HIGHER ABSOLUTE HEIGHT BECAUSE WAVE SPLASH GETS UP MUCH

HIGHER THAN IT DOES AT THE OTHER PLACE WHEN THE TIDE IS AT
A PARTICULAR LEVEL (E.G., IF THE TIDE IS AT 0.0, WAVES MIGHT BE
WASHING UP TO A HEIGHT OF +1 METER AT THE PROTECTED SPOT,
BUT ALL THE WAY UP TO +3 METERS AT THE EXPOSED SPOT). THIS
LEADS TO ORGANISMS THAT DEPEND MORE ON A MARINE
ENVIRONMENT BEING ABLE TO LIVE AT HIGHER ABSOLUTE TIDAL
HEIGHTS AT THE EXPOSED THAN AT THE PROTECTED SPOT. WE CAN
USE INDICATOR SPECIES WHOSE VERTICAL DISTRIBUTION IS
KNOWN TO DEPEND PRIMARILY ON PHYSICAL FACTORS TO
COMPARE DIFFERENT LOCATIONS IN TERMS OF RELATIVE TIDAL
HEIGHT. FOR INSTANCE, IN ONE OF THE FIGURES IN THE LECTURE
NOTES THERE IS A SHADED REGION THAT INDICATES HOW HIGH UP
THE SHORE THE ALGA ENDOCLADIA MURICATA REACHES AT EACH
OF THREE DIFFERENT LOCATIONS THAT DIFFER IN WAVE
EXPOSURE. THE TOP OF THE ENDOCLADIA ZONE CAN BE
CONSIDERED TO BE EQUIVALENT IN TERMS OF PHYSICAL FACTORS
(DESICCATION, TEMPERATURE, ETC) AT THE THREE SITES. SOTHE
TOP OF THE ENDOCLADIA ZONE WOULD BE CONSIDERED TO BE AT
THE SAME RELATIVE TIDAL HEIGHT AT THE THREE LOCATIONS,
EVEN THOUGH THE ABSOLUTE HEIGHTS ARE DIFFERENT. Why might
it be important to use relative height rather than absolute height when considering
the distribution of organisms? BECAUSE WHEN COMPARING LOCATIONS
THAT DIFFER DRAMATICALLY IN THINGS LIKE WAVE ACTION OR
EXPOSURE TO SUN (E.G., IN CAVES OR UNDER OVERHANGING
ROCKS, VERSUS OUT IN THE WIDE OPEN), ABSOLUTE HEIGHT IS NOT
VERY USEFUL. PARTICULARLY IF YOU WANT TO COMPARE THE
EXPLANATORY POWER OF ALTERNATIVE HYPOTHESES BASED ON
PHYSICAL FACTORS VERSUS, SAY, PREDATION OR COMPETITION
(BIOTIC, NOT PHYSICAL, FACTORS), ITS IMPORTANT TO THINK
ABOUT THE PHYSICAL FACTORS, AND CONSIDERING RELATIVE
TIDAL HEIGHT GIVES YOU AN EASY WAY TO DO THAT. Describe a
specific example discussed in class that illustrates how considering relative height
helps to reveal whether limpet distribution in a particular location is more likely
to be caused by predation or by physical factors. IN THE FIGURE SHOWING
THE THREE SITES WHERE OYSTERCATCHERS CAN REACH THE
LOWER PART OF THE VERTICAL SURFACE, THE SITES DIFFER BOTH
IN ABSOLUTE AND IN RELATIVE TIDAL HEIGHT. THE ONE ON THE
LEFT IS LOW BOTH ABSOLUTELY (LOOK AT ITS POSITION ON THE
RIGHT HAND Y AXIS) AND RELATIVELY (ENDOCLADIA EXTENDS
PART WAY UP THE VERTICAL SURFACE, INDICATING THAT MARINE
CONDITIONS REACH WELL UP INTO THE PART OF THE ROCK THAT
WERE INTERESTED INTHAT VERTICAL SURFACE ABOVE WHERE
THE OYSTERCATCHERS CAN WALK). THE RIGHT HAND PANEL IS AT
THE OTHER EXTREME..ITS VERY HIGH ON AN ABSOLUTE BASIS
AND ITS ALSO RELATIVELY HIGH, AS INDICATED BY THE FACT

THAT ENDOCLADIA DOESNT EVEN GET ANYWHERE NEAR THE

BOTTOM OF THE VERTICAL SURFACE WHERE THE
OYSTERCATCHERS WERE FORAGING. THE MIDDLE PANEL IS IN
BETWEEN. COMPARING LOCATIONS LIKE THESE HELP US TO BE
CONFIDENT THAT PREDATION, AND NOT ONLY PHYSICAL FACTORS,
IS CONTRIBUTING TO THE DISTRIBUTION OF THE LIMPETS.
CLEARLY, DIGITALIS CAN BE FOUND AT A VARIETY OF RELATIVE
TIDAL HEIGHTS ABOVE THE TOP OF THE ENDOCLADIA ZONE
(COMPARE THE MIDDLE AND THE RIGHT HAND PANEL). WHAT IS
CONSISTENT IS THAT ACROSS A RANGE OF RELATIVE TIDAL
HEIGHTS, LARGE DIGITALIS ARE FOUND PRIMARILY OUT OF REACH
OF OYSTERCATCHERS. WE WOULD BE MUCH LESS CONFIDENT OF
THIS RESULT IF WE USED ONLY ABSOLUTE TIDAL HEIGHT, OR
QUANTIFIED LIMPETS ONLY IN PLACES THAT WERE ALL AT
IDENTICAL RELATIVE HEIGHTS.
25. What are some key ways in which behavior of Lottia digitalis and Maclintockia
scabra differ? DIGITALIS DOES NOT HOME, AND SHOWS UPWARDMIGRATORY BEHAVIOR CAUSED BY UPWARD MOVEMENT DURING
WINTER FOLLOWED BY LESS-PRONOUNCED DOWNWARD
MIGRATION DURING SUMMER, WITH THE EFFECT THAT INDIVIDUAL
LIMPETS END UP A BIT HIGHER ON THE SHORE EACH YEAR AS THEY
GROW. THIS LEADS TO THE LARGEST DIGITALIS BEING FOUND VERY
HIGH ON THE SHORE. SCABRA HOMES VERY ACCURATELY TO A
SPECIFIC SITE AND THEY APPEAR TO REMAIN FAITHFUL TO THIS
SITE FOR A VERY LONG TIME, DEVELOPING A CLOSE FIT OF THEIR
SHELL MARGIN TO ROCK IRREGULARITIES WHEN LIVING ON HARD
ROUGH SURFACES LIKE GRANITE, AND ETCHING AN INDENTED
SCAR INTO THE ROCK SURFACE WHEN LIVING ON SMOOTH SOFT
ROCKS LIKE SANDSTONE.
26. Use an optimal foraging kind of argument to discuss the factors affecting
oystercatchers preferences for some limpets over others. In composing your
answer you should think about a number of different factors, such as energy
required to obtain the prey, energy gained from consuming the prey, time
involved in locating the prey, time involved in handling the prey once found and
attacked, efficiency with which prey is processed in the gut, etc. THERE ARE
MANY FACTORS ONE COULD INCLUDE HERE. SIZE, EASE OF
REMOVAL FROM THE ROCK, EASE OF SEPARATING UNDIGESTABLE
SHELL PARTS FROM THE LIMPET BODY, ABUNDANCE, EASE OF
ACCESS. SMALL LIMPETS MIGHT BE ABUNDANT AND THEREFORE
EASY TO FIND, AND EASY TO REMOVE FROM THE ROCK BUT HARD
TO SEPARATE SHELL FROM BODY AND OF LOW OVERALL PAYOFF
PER LIMPET. LIMPETS LIKE SCABRA MAY BE FAVORED BECAUSE OF
HIGH ABUNDANCE AND ACCESSIBILITY BUT DISFAVORED BECAUSE
THEY ARE HARD OR IMPOSSIBLE TO REMOVE FROM THE ROCK, AND
BECAUSE IF REMOVED THE SHELL IS BROKEN SO ITS HARD TO
SEPARATE DIGESTIBLE AND UNDEGESTIBLE COMPONENTS SO THE

EFFICIENCY OF PROCESSING IS LOW. ALSO, ACTUAL ENERGY

REQUIRED TO REMOVE AND HANDLE THEM WOULD BE HIGH,
MAKING THE NET PAYOFF IF EATEN RELATIVELY LOW. LIMPETS
LIKE DIGITALIS MAY BE FAVORED BECAUSE THEY ARE EASY TO
REMOVE AND HANDLE, BUT DISFAVORED BECAUSE THEY ARE
SCARCE IN ACCESSIBLE AREAS. IT WOULD BE POSSIBLE TO CREATE
MODELS THAT EVALUATE THE RELATIVE BENEFITS OF FORAGING
ON DIFFERENT LIMPETS USING THESE DIFFERENT PARAMETERS,
BUTYOU DONT NEED TO DO THAT HERE.
27. Oystercatchers tend to eat very few Maclintockia scabra on granite substrata at
Pacific Grove, and to eat large numbers of Lottia digitalis and other limpets there
despite the fact that those limpets are much less prevalent than scabra in areas
accessible to the birds. However, oystercatchers are relatively uncommon there,
so the magnitude of the predation pressure is relatively low. Use an optimal
foraging argument to predict what would happen to oystercatcher prey selectivity,
as well as to limpet distribution and abundance, if the density of foraging
oystercatchers were to increase dramatically around Pacific Grove. ONE
POSSIBLE OUTCOME WOULD BE THAT IF THE DENSITY OF
OYSTERCATCHERS BECAME HIGH ENOUGH, THE AVAILABILITY OF
PREFERRED PREY LIKE DIGITALIS IN AREAS THAT ARE IN REACH OF
OYSTERCATCHERS COULD GET SO LOW THAT OYSTERCATCHERS
WOULD START TO EAT SCABRA. IF SCABRA ARE ACTUALLY
IMPOSSIBLE (AS OPPOSED TO JUST DIFFICULT) FOR
OYSTERCATCHERS TO REMOVE AND/OR HANDLE, THEN THE
SWITCH TO SCABRA WOULD NOT HAPPEN. BUT IF THE REASON
SCABRA ARE NEGLECTED MAINLY HAS TO DO WITH THEIR BEING
RELATIVELY UNPROFITABLE TO EAT AS COMPARED WITH OTHER
LIMPETS SUCH AS DIGITALIS, THEN THEIR ABSENCE IN THE
OYSTERCATCHER DIET IS DEPENDENT ON OTHER LIMPETS BEING
AVAILABLE IN SUFFICIENT NUMBERS TO KEEP OYSTERCATCHERS
FOCUSED ON THOSE OTHER SPECIES. IF OYSTERCATCHER
ABUNDANCE GOT HIGH ENOUGH, WE MIGHT EXPECT TO SEE ALL
LIMPETS OF ACCEPTABLE SIZE RESTRICTED TO AREAS OUT OF
REACH OF OYSTERCATCHERS. AND UNDER THIS SCENARIO,
SCABRA MIGHT BECOME EXTREMELY SCARCE, SINCE DIGITALIS
WOULD BE EXPECTED TO OUTCOMPETE SCABRA IN THE SAFE
PLACES.
28. Some rock types (e.g., greenstone, such as is prevalent in some parts of the Big
Sur coast of central California) are extremely smooth and very hard too smooth
to provide physical structure for the shell to interact with and too hard for scabra
to etch home scars into them. Based on what you know of oystercatcher prey
preference and limpet distribution on granite and sandstone, and of limpet
behavior on these different types of rocks, how would you expect oystercatcher
prey selectivity and limpet distribution and abundance to differ on smooth hard
rocks like greenstone from what has been observed on granite and sandstone
shores? I WOULD EXPECT LIMPETS OF ALL SPECIES BIG ENOUGH TO

BE OF INTEREST TO OYSTERCATCHERS (I.E., ANY LIMPET BIGGER

THAN 10MM SHELL LENGTH) TO BE SCARCE ON ANY SURFACE
ACCESSIBLE TO OYSTERCATCHERS. FURTHER, SCABRA MAY BE
UNIFORMLY SCARCE IN THIS TYPE OF AREA (LIKE IN AN AREA WITH
VERY HIGH OYSTERCATCHER ABUNDANCE, AS IN QUESTION 16
ABOVE). AROUND PACIFIC GROVE, THE REASON SCABRA ARE ABLE
TO BE SO ABUNDANT IS THAT HOMING PROVIDES SCABRA A
REFUGE FROM OYSTERCATCHER PREDATION IN AREAS THAT ARE
UNAVAIALBLE TO SUPERIOR COMPETITORS LIKE DIGITALIS.
SCABRA WOULD HAVE NO SUCH REFUGE FROM COMPETITION IN AN
AREA SUCH AS DESCRIBED HERE. Explain your reasoning, and include a
discussion of what limpet species and sizes you would expect to be in the
oystercatchers diets, and where you would expect to find limpets of different
species and sizes living in this type of area. PARTLY ANSWERED ABOVE. I
WOULD EXPECT OYSTERCATCHER DIET TO INCLUDE ALL SPECIES
MORE OR LESS IN PROPORTION TO THEIR ABUNDANCE, AND
PROBABLY SIZE PREFERENCES WOULD BE SIMILAR TO WHAT HAS
BEEN MEASURED ELSEWHERE, THOUGH IF THEY HAVE DEPRESSED
NUMBERS OF LARGER LIMPETS SUFFICIENTLY THEY MAY BE
FORCED TO ACCEPT MORE SMALL LIMPETS (RIGHT AROUND 10MM
SHELL LENGTH) THAN ELSEWHERE. NOTE THAT THE LOWER SIZE
LIMIT PROBABLY HAS PARTLY TO DO WITH PHYSICAL
CONSTRAINTS IMPOSED BY THE OYSTERCATCHER BILLTHE TIP IS
KIND OF CHISEL-SHAPED, SO ONLY LIMPETS LARGER THAN ABOUT
10MM LENGTH ARE ABLE TO BE HANDLED EFFICIENTLY AND
REMOVED FROM THEIR SHELLS ONCE POPPED OFF THE ROCK.