CHAPTER 55

BEHAVIORAL
ECOLOGY

Prepared by
Brenda Leady, University of Toledo

Copyright (c) The McGraw-Hill Compa 1

nies, Inc. Permission required for repro
 Behavior is the observable response of
organisms to internal or external stimuli
 Behavioral ecology studies how behavior
contributes to the differential survival and
reproduction of organisms
 Ethology focused on the physiological
mechanisms of behavior
 Proximate causes – change in day length
 Behavioral ecologists focus on the adaptive
significance of a behavior
 Ultimate causes – effect on reproductive success

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 Some Behavior Results from Simple
Genetic Influences
 W.C. Rothenbuhler’s 1964 work showed a genetic basis to behavior
in honeybees
 Hygienic bees removed diseased larvae using 2 distinct maneuvers –
uncapping cells and removing larvae
 Strains that were not hygienic did not do this
 u – uncapping, U – did not uncap
 r – larval removal, R – no removal
 uurr – hygienic, UURR – not hygienic
 UuRr – not hygienic, uuRr – uncap cells but larva not removed, Uurr – remove
larva if cells uncapped
 Jennifer Brown in 1996 found fosB gene for nurturing in mice
 Normal mice clean pups, nurse them, crouch over them
 Mutant mice lacking fosB do not do these behaviors and pups die
 Single gene can have dramatic impact on behavior
Impact of genetics and learning on behavior

 Fixed action patterns (FAP)

 Innate or genetically programmed behavior
 Once initiated, will continue until completed
 Egg-rolling response in geese
 Improves fitness because it increases survival of
young
 Sign stimulus – initiates behavior
 Egg out of nest
 Male sticklebacks attack red ventral surface while ignoring
realistic fish model lacking red underside

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 Learning – modify behavior based on
previous behavior
 Habituation – simplest form of learning
 Organism ignores repeated stimulus
 Form of nonassociative learning – decrease in
response to stimulus due to repetition

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 Associative learning - association develops
between stimulus and response
 2 main types
1. Classical conditioning – involuntary response
becomes associated positively or negatively with a
stimulus that did not originally elicit the response
 Pavlov’s dog salivates when the bell rings
 Food is the unconditioned stimulus
 Bell is the conditioned stimulus
 Salivation in response to food is the unconditioned response
 Salivation in response to the bell is conditioned response

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2. Operant conditioning – animal’s behavior
reinforced by a consequence (reward or
punishment)
 Skinner box where rat bumps into a lever and
gets food
 Associate lever with food
 Also called trial-and-error learning
 Birds will learn to avoid bad tasting butterflies

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(a) Blue Jay eating monarch (b) Blue Jay vomiting

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 Cognitive learning – ability to solve problems
with conscious thought and without direct
environmental feedback
 Chimpanzees stack boxes to reach banana
 Ravens retrieve meat by pulling up a string

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 Behavior is often a mix of innate and learned
 Birds are genetically programmed to learn but they
will sing the correct song only if the correct songs are
heard

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 Critical period – time when many animals
develop species-specific patterns of
behavior
 Imprinting – goslings follow the first moving
thing as “mother”
 Innate behavior is the ability to imprint
 Factors in the environment are the stimulus to
which imprinting is directed
 Migration– experienced birds can correct for
displacement (complex navigational skill)
while young, inexperienced birds cannot
correct
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Local and long-range migration
 Local movements
 Movements to find food, water, nesting site
 Kinesis – movement in response to stimulus
but not directed toward or away from source
 Taxis – more directed movement
 Positive phototaxis – toward light
 Negative phototaxis – away from light

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Tinbergen’s Experiments Show Digger Wasps Use
Landmarks to Find Their Nests

 Digger wasp females provide honeybees for larvae to

feed on
 Each larva needs 5 to 6 bees which must be brought
individually to nest
 Tinbergen hypothesized wasps found nest by creating
mental map of area
 First, experimentally adjusted landmarks
 Wasps flew to moved pinecones
 Second, tested if it was sight or smell of pinecones
 Found it was sight of cones used as map
 Migration – long-
range seasonal
movement
generally linked to
seasonal
availability of food
 Bird, mammal,
and insect
examples
 3 mechanisms to
find their way

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1. Piloting – animal moves from one familiar
landmark to the next
 Features of the coastline, for example
2. Orientation – ability to follow compass bearing
and travel in straight line – cannot adjust for
course
3. Navigation – follow compass bearings but
also set or adjust path
 Adult starlings can adjust flight path when
transported and released (juveniles cannot)

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 Many species use a combination of navigational
reference points and an internal clock
 Pigeons integrate internal clock with position of the sun –
for every hour their internal clocks were shifted, their
orientation shifted 15°
 Not all migrations well understood
 Green sea turtles migrate to Ascension Island to lay eggs
 Homing pigeons can be transported to sites they have
never been to and fly directly home

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Foraging
 Optimality theory predicts an animal
should behave in a way that maximizes
benefits of a behavior minus its costs
 Optimal foraging proposes that an animal
seeks to obtain the most energy possible
with the least expenditure of energy
 Themore net energy gained, the greater the
reproductive success

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 Shore crabs will eat
different sized
mussels
 Prefer intermediate
mussels with highest
rate of energy return
 Larger mussels yield
more energy but take
longer to open
 Smaller mussels are
easier to open but
yield less energy

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 Defending a territory has costs and
benefits
 Territory– fixed area in which individual or
group excludes others
 Optimize territory size based on costs and
benefits
 Benefit is exclusive access to resource- food,
mates, nesting sites
 Costly to defend

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 Golden-winged
sunbird
 Saved 780 calories
a day in reduced
foraging activity
 Spent 728 calories
in defending the
territory
 Net gain of 52
calories a day
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 Cheetahs need
large territories
relative to body size
to hunt successfully
 Gannet territory
size determined by
how far bird can
reach to peck its
neighbor without
leaving nest

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Communication
 Use of specifically designed signals or
displays to modify the behavior of others
 Chemical
 Auditory
 Visual
 Tactile

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 Chemical communication
 Common among canines and felines
 Scent trails laid by social insects
 Pheromones produced by female moths to attract
males
 Queen bee releases pheromones to suppress
reproductive system of workers
 Auditory communication
 Sounds travel farther in air
 Air at dawn and dusk less turbulent
 Many males use auditory communication to attract
females
 Sound production can also lure predators

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 Visual communication
 Competition among males for most impressive
displays leads to elaborate coloration and
extensive ornamentation
 Male fireflies flash species specific number and
duration of flashes
 Predator uses flashes to lure males in to eat them
 Tactile communication
 Used to establish bonds between group members
 Round dance or waggle dance of honeybee scout
conveys food location

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Living in groups
 Much of animal behavior directed at other
animals
 Some of the more complex behavior
occurs in groups like flocks or herds
 Group living can reduce predation through
 Increased vigilance
 Protection in numbers

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 Increased vigilance
 Many eyes hypothesis – by living in groups,
individuals may decrease the amount of time
scanning for predators and increase time
feeding
 If each pigeon occasionally looks up to scan
for a hawk, the bigger the group, the more
likely that one bird will spot a hawk early
enough for the flock to take flight

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 Protection in numbers
 Typically, predators take one prey item per attack
 In a large group, chances of being that prey item are
reduced
 “Selfish herd” – each individual can minimize the
danger to itself by choosing the location that is
closest to the center of the group
 Group size may be the result of trade offs between
the benefits of group living and costs like grooming
and altruism

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Altruism
 Behavior that appears to benefit others at
a cost to oneself
 Most altruistic acts serve to benefit the
individual’s close relatives

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 Individual selfish behavior is more likely
 Group selection – group containing altruists would
have a survival advantage over group composed of
selfish individuals
 Individual selection more likely because…
 Mutant individuals that use resources have an
advantage over those that conserve resources
 Selfish individuals can immigrate from other areas

 For group selection to work, groups must die faster than

others – individuals die more often than groups

 Group selection assumes that individuals can predict

food availability to conserve resources as needed – little

evidence that they can

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 Example of selfish behavior
 Male Hanuman langurs kill infants when they take
over groups of females from other males
 When not nursing, females become sexually receptive
sooner, so a male can father offspring sooner
 Infanticide ensures that the male will father more
offspring
 Genes governing this trait spread by natural selection

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 Kin selection
 Coefficient of relatedness – probability that any 2
individuals will share a copy of a particular gene is a
quantity r
 An organism can not only pass on its genes by having
offspring but also by ensuring that relatives survive
 Inclusive fitness designates the total number of copies
of genes passed on through one’s relatives or as
one’s own offspring
 Kin selection – behavior that lowers an individual’s
own fitness but enhances the reproductive success of
relatives

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 Hamilton’s rule
 Altruistic gene favored by natural selection
when

rB>C
r is the coefficient of relatedness of donor
(altruist) to recipient
 B is benefit to recipient
 C is cost incurred by donor

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 Datana caterpillars example
 Brightly colored and assume specific pose
when threatened
 Predator has to eat one to learn to avoid them
 Death of individual in group of related
caterpillars benefits siblings
 r =0.5, B=50, and C=1, then 25(0.5x50)>1 so
genes will spread

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 Altruism in social insects due to genetics and
lifestyle
 Most extreme form of altruism is sterile castes in
social insects
 Eusociality – workers (females) help queen raise
offspring
 Haplodiploidy – females are diploid, males are haploid,

females are more related to their sisters (0.75) than they

would be to their own offspring (0.5)
 Existence of eusocial mammals predicted based on
lifestyle
 Naked mole rats have a queen who suppresses
reproduction in other females

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 Reciprocal altruism
 Cost to the altruist offset by likelihood of a
return benefit
 Female vampire bats will share food
 Unrelated females are more likely to share
food with those that had recently shared with
them

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Mating systems
 Natural selection favors production of the rarer
sex so that the sex ratio is kept balanced at 1:1
 Monogamy – each individual mates exclusively
with one partner
 Polygamy – individuals mate with more than one
partner
 Polygyny – one male mates with many females
 Polyandry – one female mates with many males

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 Sexual selection
 Promotes traits that will increase an
organism's mating success
 2 forms
 Intersexual – member of one sex chooses mate
based on particular characteristics
 Intrasexual – members of one sex compete over
partners with the winner performing most of the
matings

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 Female mate choice
 Female hangingflies demand a nuptial gift –
allows female to produce more eggs and
allows male to copulate longer
 Female sticklebacks prefer males that shake
more during courtship as evidence that he will
be a better parent
 Choices based on plumage color or courtship
displays – widowbird with experimentally
lengthened tails attracted more females and
fathered more clutches

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 Mate competition between individuals
 In many species, females do not actively
choose between mates
 Instead they mate with competitively superior
males
 Dominance determined by fighting or
ritualized sparring
 Male-male competition produces males
substantially larger than females
 Small males can still father offspring by
intercepting females

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 Monogamy
 One male mates with one female
 Males and females generally similar in body size and
appearance
 Mate-guarding hypothesis – males stay with a female
to protect her from being fertilized by other males
 Male assistance hypothesis – males remain with
females to help them rear offspring – he would have
few surviving offspring if he did not
 Female-enforced monogamy hypothesis– female
interferes with male attracting other females

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 Polygyny
 One male mates with more than one female
 Females mate with only one male
 Associated with uniparental care of young
 Males contribute little to raising young
 Sexual dimorphism typical
 Types
 Resource based polygyny – patchy distribution of resource
and female visits for resource
 Harem mating structure – females naturally congregate and

male controls area

 Communal courting – males display in lek, females mate

after males display

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 Polyandry
 One female mates with several males
 Rarer
 Female is larger of the sexes
 Female spotted sandpiper reproductive
success limited only by the number of males
she can find to incubate her eggs
 Male pipefish have brood pouches and female
produces enough eggs for 2 male brood
pouches if she can find another male

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