Chapter 56

5th: 12891317
4th: 1334-1363
Animal Behavior

• Instinctive & Learned behaviours

• Neurophysiological & Endocrine
control of behaviour
• Migration & wayfinding
• Habitat Selection & Territoriality
• Evolution of communication
• Evolution of reproductive behaviour
& mating systems
• Evolution of social behaviour

Russell, Biology: The Dynamic Science, 5th edition. © 2021 Cengage. All Rights Reserved. May not be scanned, copied
or duplicated, or posted to a publicly accessible website, in whole or in part.
Learning Objectives
• Identify general categories of behaviors that are
typically instinctive and those that are typically
learned.
• Compare and contrast the three mechanisms
responsible for wayfinding by migrating animals.

• Cite evidence to support the hypothesis that variations in the

availability of food is responsible for the seasonal migration of
many animals
• Observe the interaction between two individuals of the same
species and identify the types of signals they use for
communication.
Learning Objectives (3 of 3)
• Explain how differences in the reproductive
strategies of males and females can foster sexual
selection for certain characteristics in males.
Animal Behavior and Ethology
• The study of animal behavior involves discovering how
animals respond to specific stimuli and why they respond in
predictable and characteristic ways.
• European researchers developed a comprehensive approach
to animal behavior studies (ethology), which focuses on
how animals behave in their natural environments.
• They analyzed how evolutionary processes shape
inherited behaviors and how animals respond to specific
stimuli.
Behavioural ecology, begins with the assumption that
behaviour is subject to natural selection.
Behaviour has:
i) a genetic basis
ii) lead to increased survival & nr of offspring

THUS, behaviours of animals we observe today must

have adaptive values (such as territory, reproductive
strategies, altruism)
Behavioural repertoire includes a variety of actions
and responses.

The study of Animal Behaviour involves discovering

how animals respond to specific stimuli and why
they respond in predictable and characteristic
ways.
The discipline of ethology developed in mid 70’s with
Konrad Lorenz, Niko Tinbergen & Karl von Frisch

It focuses on how evolutionary processes shape inherited

behaviours and the ways that animals respond to specific
stimulus.
Animal Behavior and Ethology
• Niko Tinbergen identified four basic questions that
any broad study of animal behavior should address:
1. What mechanisms trigger specific behavioral
responses?
2. How does a behavior develop as an animal
matures?
3. What is the behavior’s function and how does it
increase an animal’s chances of surviving and
reproducing?
4. How did the behavior evolve?
Proximate Causes of Behavior
• Advances in neuroscience — the integrated study
of the structure, function, and development of the
nervous system —now allow researchers to explore
Tinbergen’s first and second questions about the
proximate causes of behavior:
• how genetic, cellular, physiological, and
anatomical mechanisms underlie an animal’s
ability to detect and respond to stimuli in
species-specific ways.
Ultimate Causes of Behavior
• Comparable advances in genetic analysis and
evolutionary theory enable scientists to address
Tinbergen’s third and fourth questions about the
ultimate causes of behavior:
• why behaviors have adaptive value;
• why the genetic and physiological mechanisms
that underlie specific behaviors are subject to
microevolutionary change.
56.1 Instinctive and Learned
Behaviors
• Instinctive behavior is genetically programmed – it
appears in complete and functional form the first
time it is used.
• Learned behavior is dependent upon having a
particular kind of experience during development.
• No behavior is determined entirely by genetics or
entirely by environmental factors – behaviors
develop through complex gene–environment
interactions.
Song of the White-Crowned
Sparrow
• Observation: Adult male white-crowned sparrows sing a song that no
other species sings.
• Hypotheses:
• If the song is instinctive, isolated male nestlings should be able to
sing their species’ song when they mature;
• If the song is learned, isolated male nestlings should not sing
“properly” when they become adults.
• Experiment: Peter Marler reared white-crowned sparrows individually in
soundproof cages in his laboratory.
Song of the White-Crowned
Sparrow
• Method: Some 10 to 50 day-old chicks listened to
recordings of a male white-crowned sparrow’s
song, others did not.
• Results: When juvenile males in both groups first
started to vocalize at about 150 days, they
produced only whistles and twitters.
• At about 200 days of age, young males that had
listened to recordings were producing the song
they had heard.
• Males in the group that had not heard recordings
never sang the way wild males do.
Song of the White-Crowned
Sparrow
• Conclusion: learning is essential for a young male
white-crowned sparrow to acquire the full song of
its species.
• Although early researchers classified behaviors as
either instinctive or learned, we now know that
most behaviors include both instinctive and learned
components.
• Some behaviors have a strong instinctive basis;
others are mostly learned.
Instinctive Behaviors
Behavior Description
Instinctive behaviors
(strong genetic basis)
Fixed action patterns Stereotyped actions, often
initiated by a sign stimulus
Feeding behaviors Innate food preferences
and hunting tactics
Defensive behaviors Responses to predators
Reproductive behaviors Mating habits and parental
care activities

Russell, Biology: The Dynamic Science, 5th edition. © 2021 Cengage. All Rights Reserved. May not be
scanned, copied or duplicated, or posted to a publicly accessible website, in whole or in part.
Instinctive Behaviors
• Instinctive behaviors can be performed without
prior experience.
• We assume that they have a strong genetic basis
and that natural selection has preserved them as
adaptive behaviors:
• Examples: Feeding behaviors, defensive
responses, mating behaviors, and parental care
activities.
Stereotyped Behaviors
• Many instinctive behaviors are highly stereotyped
– when triggered by a specific cue, they are
performed over and over in almost exactly the
same way.
• Such behaviors are called fixed action patterns,
and the simple cues that trigger them are called
sign stimuli:
• Example: In herring gull chicks, begging for food
(the fixed action pattern) is triggered when they
see a red spot on the lower bill of an adult (the
sign stimulus).
Stereotyped Behaviors
• Natural selection has molded the behavior of some
parasitic species to exploit the relationship
between sign stimuli and fixed action patterns:
• Example: Birds that are brood parasites lay their
eggs in the nests of other species – exaggerated
sign stimuli from the chick (opening its mouth,
bobbing its head, and calling vigorously) elicit
feeding behavior by the foster parents.
Do Animals Have Emotions?
• Body language of animals can be interpreted to suggest that
they have feelings. Ex wolves, elephant.

• Today scientists believe that they have sufficient data to

suggest that many vertebrates have feelings such as: Fear,
joy, embarrassment, jealousy, love, anger, sadness, fear,
grief
Animals can feel love, but perhaps not to the degree that
humans can.

It is unlikely that emotions 1st appeared in humans with

no evolutionary homologies in animals.
Researchers have found a high level of dopamine in
the brain when rats play, and the dopamine level
increases even when rats anticipate the opportunity to
play.

Studying animal emotions may best be left to field

research; laboratory animals may be too stressed to
provide convincing data on emotions.
Genes and Behavior
• Although single genes can influence behavior, they
do not control complex behavior patterns directly.
• The alleles present affect the kinds of enzymes that
cells can produce, influencing the biochemical
pathways involved in development of an animal’s
nervous system.
• Resulting neurological differences can translate into
a behavioral difference between individuals that
have certain alleles and those that do not.
Learned Behaviors
• Learning is a process in which experiences change
an animal’s behavioral responses.
• Different types of learning occur under different
environmental circumstances.
Learned Behaviors
Behavior Description
Learned behaviors (strong
experiential basis)

Imprinting Affinity for caretaker species,

developed during critical period

Classical conditioning Association between

phenomena that are usually
unrelated
Operant conditioning Trial-and-error learning

Cognition Insight learning in novel

situations
Habituation Loss of responsiveness
Russell, Biology: The Dynamic Science, 5th edition. © 2021 Cengage. All Rights Reserved. May not be
scanned, copied or duplicated, or posted to a publicly accessible website, in whole or in part.
Imprinting
• In imprinting, animals learn the identity of a caretaker
or the key features of a suitable mate during a
restricted stage of development (critical period) early
in life:
• Example: Newly hatched geese imprint on their
mother’s appearance and identity, staying near her
for months.
• At sexual maturity, geese mate with another goose
because it exhibits the visual and behavioral stimuli on
which they imprinted when young.
Imprinting (=form of learning)

KONRAD LOREZ
Imitation of behaviour observed during sensitive
period (can be 2-3 days after birth in the case of birds)
KL studied chicks, ducks + goslings, follow any moving
object after birth
• Sensitive period = Period of time in which a particular
behaviour develops
• Ex a chick imprinted on a red ball follows it around and
chirp the moment the ball is out of sight
Classical Conditioning

• Ivan Pavlov experimented with classical conditioning,

a type of learning in which animals develop a mental
association between two unrelated phenomena.
• Dogs salivate when they eat – food is an
unconditioned stimulus because dogs respond to it
instinctively, no learning is required for the stimulus to
elicit the response (salivation).
Classical conditioning suggests that an organism
can be trained (conditioned) to associate any
response with any stimulus.

Unconditioned responses are those that occur

naturally (when salivation follows the
presentation of food).

Conditioned responses are those that are

learned (dog that learned to salivate at the ring of
a bell)
Operant Conditioning
(reinforcement)
• In operant conditioning (trial-and-error learning)
animals learn to link a voluntary activity (an operant)
with a reward (a reinforcement):
• Example: If a rat accidentally presses a bar (the
operant) and receives food (the reinforcement), it
will learn to press the bar more frequently – as long
as it produces results.
• Laboratory rats have also learned to press bars to turn
off disturbing stimuli, such as bright lights.
Cognition
• Animals can solve a novel problem by using insight
– “thinking up” a solution rather than making a
series of trial-and-error attempts at resolving it.
• This process (cognition) implies that an animal is
aware of its circumstances, defines a specific goal,
and then uses reasoning to achieve the goal:
• Example: Ravens used cognition to figure out how to
obtain food that dangled below their perches on a
string.
Ravens were offered meat attached to a
string hanging from a branch, custom to
eating meat but had no knowledge of
how the string works.

Several hour later, one flew to branch,

grab string with its beak, pulling the
meat closer.

Sea otters “save” particular rocks that act as hard

surface to bash open clams and mussels.
Habituation
• Animals typically lose their responsiveness to
frequent stimuli that are not quickly followed by
the usual reinforcement.
• Learned loss of responsiveness (habituation) saves
time and energy of responding to stimuli that are
not important:
• Example: The sea hare responds to a touch by
retracting its delicate gills – but, if it is touched
repeatedly with no harmful consequences, it
stops retracting its gills.
Animals loses their responsiveness to frequent stimuli that
are not quickly followed by usual reinforcement.

This learned loss of responsiveness habituation save

animals the time + energy of responding to stimuli that are
no longer important.

Habituation is important in filtering the large quantities of

information received from the surrounding environment.

By habituating to less important signals, animals can focus

their attention on the more important features of the
environment.
Prairie dogs, Cynomys ludovicianus, give alarm calls when
mammals, large birds, or snakes approach.

Individual prairie dogs are particularly susceptible to becoming

food for a coyote, hawk, or rattlesnake, but collectively they are
quite well-defended, as their alarm calls facilitate escape in
burrows.

When prairie dog occurs around town located near trails

used by humans, giving alarm calls every time a person
walks by is a waste of time and energy for the group.

Habituation to humans is an important adaptation in this

context.
56.2 Neurophysiological and
Endocrine Control of Behavior
• All behavioral responses depend on the
biochemistry and structure of neurons (nerve cells).
• Although the anatomical and physiological basis for
some behaviors is present at birth, an individual’s
experiences alter cells of its nervous system in ways
that produce particular patterns of behavior:
• Example: The singing behavior of songbirds.
Neural Circuits in Songbirds
• Acoustical experience shapes singing behavior in
adult male white-crowned sparrows.
• A sparrow chick’s brain acquires and stores
information present in the songs of other males.
• When the young male starts to sing, its nervous
system matches the vocal output to the stored
memory of the song.
• When it achieves a good match, the sparrow’s brain
must “lock” on the complete song.
Neural Circuits in Songbirds
• Certain neurons in the young male’s brain are
influenced only by appropriate stimuli (acoustical
signals from individuals of its own species) and only
during the critical period.
• Neuroscientists have identified neuron clusters
(nuclei) in the brain that make song learning and
song production possible.
• In male zebra finches, a nucleus in the forebrain
responds to songs of strangers but not to songs of
established neighbors on adjacent territories.
Hormonal Changes and Behavior
• A change in concentration of a certain hormone is
often the physiological trigger that induces
important changes in an animal’s behavior as it
matures.
• In honeybees, increasing concentration of juvenile
hormone induces a change in the tasks worker bees
perform:
• Worker bees less than 15 days old care for
larvae and maintain the hive;
• Older bees leave the hive to collect nectar and
pollen.
Hormones and Reproductive
Behavior
• Territorial male African cichlid fish maintain nesting
territories on the bottom of Lake Tanganyika in East
Africa, and defend them aggressively against other
males.
• These territorial males are brightly colored and
exhibit elaborate behavioral displays that attract
females.
• Nonterritorial males are less colorful and aggressive
– they control no nesting habitat and make no
effort to court females.
Hormones and Reproductive
Behavior
• Behavioral differences between the two types of male
cichlids are caused by differences in their levels of
circulating sex hormones.
• In territorial fish, GnRH-producing neurons in the
hypothalamus are large and biochemically active – in
nonterritorial fish, they are small and inactive.
• In the absence of GnRH, testes do not produce
testosterone – the testosterone-deficient fish neither
court females with sexual displays, nor attack other
males.
56.3 Migration and Wayfinding
• Many animal species undertake a seasonal migration,
traveling from the area where they were born to a distant
location, and returning to their birth site later.
• Examples:
• Arctic terns fly in an annual roundtrip migration of up to
70,000 km;
• Gray whales and salmon swim on long-term migrations;
• Spiny lobsters walk in long lines between coral reefs and
the open ocean floor on a seasonal cycle.
Migratory behaviour has 2 causes:

1) Proximate: this is due to environmental stimuli, that

will tell the birds it is time to travel.

2) Ultimate cause is due to the possibility of reaching a

more favourable environment for survival and
reproduction.
Migration = Long-distance travel from one location to
another.
EX: Loggerhead sea turtles hatch on Florida beach and
travel across Atlantic Ocean to the Mediterranean.

Monarch butterflies fly from North America to Mexico

so they can continue breeding.
Wayfinding Mechanisms
• Wayfinding mechanisms used by animals to arrive
at their destination are grouped into three general
categories:
• Piloting
• Compass orientation
• Navigation
• Many species use a combination of these
mechanisms to guide their movements.
Piloting
• The simplest wayfinding mechanism is piloting, in
which animals use familiar landmarks to guide their
journey:
• Example: Gray whales migrate from Alaska to Baja
California and back using visual cues provided by the
Pacific coastline.
• Animals that do migrate use specific landmarks to
identify nest sites or places where they have
stashed food:
• Example: Experiments show that digger wasps respond
to the geometry of landmarks around their nests.
Compass Orientation
• In compass orientation, animals move in a
particular direction over a specific distance or time.
• Some migratory animals use polarized light or
Earth’s magnetic field as a compass.
• Some day-flying migratory birds orient themselves
using the sun’s position in conjunction with an
internal biological clock.
• Some birds that migrate at night use the positions
of stars to determine their direction (Example:
indigo bunting).
Navigation
• The most complex wayfinding mechanism is
navigation, in which an animal moves toward a
specific destination using both a compass and a
“mental map” of where it is in relation to the
destination.
• True navigation has been documented in only a few
animal species, such as the homing pigeon, which
probably uses the sun’s position as a compass and
olfactory cues as a map.
Evolution of Migratory Behavior
• Migratory behavior entails obvious costs, such as
the time and energy devoted to the journey and
the risk of death from exhaustion or predator
attack.
• Benefits of migration include seasonal changes in
food supplies and avoiding the cold of winter:
• Example: Monarch butterflies eat milkweed in North
America during spring and summer, then migrate 4,000
km to winter in central Mexico – the round trip takes
two to five generations.
56.4 Habitat Selection and
Territoriality
• An animal’s choice of habitat is critically important
because the habitat provides food, shelter, nesting
sites, and the other organisms with which it
interacts.
• If an animal chooses a habitat that does not
provide appropriate resources, it will not survive
and reproduce.
• If an animal doesn’t find the food or other
resources it needs, or if other individuals have
already depleted those resources, it may move to
another habitat patch.
Finding Habitats
• Some animals find suitable habitats by kinesis – a
change in rate of movement or frequency of
turning movements in response to environmental
stimuli:
• Example: Wood lice move in response to dry conditions.
• Other animals exhibit a taxis response that is
directed either toward or away from a specific
stimulus:
• Example: Cockroaches exhibit negative phototaxis – they
actively avoid light and seek darkness.
Genetic Influence on Habitat
Selection
• Biologists assume that habitat selection is adaptive
and has a genetic basis shaped by natural selection.
• Examples:
• Some animals instinctively select habitats where they
are well camouflaged;
• Many insects have a genetically determined preference
for plants they eat during their larval stage;
• Blue tits and coal tits instinctively choose different trees
where each species feeds most successfully.
Learning Influences Habitat
Selection
• Habitat preferences can be molded by experiences.
• Example: Habitat choice in tadpoles:
• Tadpoles of red-legged frogs live in habitats cluttered
with sticks, algae, and plant stems – they prefer striped
backgrounds to plain ones;
• Tadpoles of cascade frogs live over gravel bottoms –
they prefer plain substrates over striped ones;
• When red-legged frogs are reared over plain substrates
and cascade frogs over striped substrates, they no
longer exhibit preferences for their usual substrates.
Territoriality
• Some animals defend a territory from other
members of their species, retaining control of the
resources it contains.
• In many organisms, territorial behavior occurs only
during the breeding season, triggered by elevated
levels of hormones.
• Animals establish and defend territories only if
some critical resource is in short supply – the
resource must be fixed in space so that the area
around it can be defended.
Territoriality

• Territorial defense is a costly activity – patrolling

territory borders, performing displays hundreds of
times per day, and chasing intruders take time and
energy.
• Territorial displays also increase an animal’s
likelihood of being injured or captured by a
predator.
• Benefits of maintaining a territory include access to
nesting sites, food supplies, and refuges from
predators.
56.5 The Evolution of
Communication
• Territorial behavior provides a specific example of
communication – the conveyance of information to
other individuals.
• All communication systems involve an interaction
between a signaler that transmits information, and
a signal receiver that intercepts the information
and makes a behavioral response.
• Natural selection has adjusted the ability of
signalers to transmit information and the ability of
receivers to get the message.
Animal Signals
• Acoustical signals (e.g., bird songs)
• A signaler produces a sound heard by a signal receiver.
• Sounds reach distant receivers, even at night and in
cluttered environments where vision is less effective.
Language = ultimate auditory communication.

Only humans can produce many different sounds and

assemble them in many different ways.

Nonhuman primates are limited to about 40 distinct

vocalizations with limited meaning.

Ex. vervet monkey's alarm calls

Auditory communication (acoustical signals) has
advantages over other kinds of communication.

i) It is faster than chemical communication

ii) It is effective both day and night
iii) Can be modified by loudness, pattern, repetition, and
duration.
Male crickets have calls for reproduction.

Birds have various songs for distress, courting, and

marking territories.

Whale songs have six basic themes for sexual and

group identification.

Bottlenose dolphins have one of the most complex

languages in animal kingdom.
Visual signals (e.g., facial expressions and body language)
In many animals, visual signals are ritualized –
exaggerated into an easily recognized visual display.
Fireflies and certain fishes send bioluminescent
signals in dark environments.
Visual communication, allows animals to signal others
of their own intentions, without the need to provide
chemical or auditory messages.

Visual signals are most often used:

i) By species that are active during the day
ii) In contests between males who make use of
threat postures

Ex. Male baboons, establish his dominance, thus keeping

peace in the troop
Animal Signals

• Chemical signals (e.g., scent marking by male dogs)

• Carry olfactory messages.
• Mammals and insects use pheromones, chemical
signals that influence the behavior of members of
the same species.
• Pheromones attract mates – a male silkworm moth can
recognize a single molecule of bombykol, a pheromone
produced by female silkworms.
• Worker ants use different pheromones to recruit fellow
workers to fight or to collect food.
Animal Signals

• Tactile signals (e.g., grooming in macaws)

• Touch signals operate only over very short distances.
• For social animals, they play a significant role in the
development of friendly bonds between individuals.
• Gull chicks peck at the parentʼs beak in order to
induce the parent to feed them
• A male leopard nuzzles the female’s neck to calm her
and to stimulate her willingness to mate.
• In primates, grooming helps cement social bonds in the
group
Communication in Honeybees
• The dance of the honeybee involves tactile, acoustical,
and chemical communication.
• When a honeybee discovers food, it returns to its
colony and performs a complex dance that conveys
information about the distance and direction of the
food source.
• When the food source is nearby, the bee performs a
“round dance” and regurgitates a food sample that
serves as a chemical cue for other workers to search for
that type of food.
• If the food source is more distant, the forager performs a
“waggle dance” – moving in alternating half-circles and
straight lines, and producing a brief buzzing sound.
• The angle of the straight run relative to vertical on the
honeycomb indicates the direction of the food source
relative to the position of the sun.
• The duration of the waggles and buzzes that the bee makes
on the straight run indicates the distance to the food.
•Electrical signals (e.g., electric eel)
Some fishes have electric organs that
release charges of variable intensity,
duration, and frequency, allowing
substantial modulation of the message.
Electrical discharges can signal threats,
submission, or readiness to breed.
Evolutionary Analysis

• Signal receivers often respond in predictable ways:

• A male white-crowned sparrow avoids entering a neighboring
territory if it hears the song of a resident male.
• Young male baboons often retreat without a fight when they see
an older male’s visual threat display.
• Biologists hypothesize that avoiding conflict may allow the
male to contribute more offspring to the next generation
by not wasting time and energy on a battle he is likely to
lose.
• Evolutionary analysis helps explain the call of ravens
that scavenge carcasses in northern forests during
winter.
• Paired, territory-owning adult ravens do not call loudly
when they find a carcasses, but feed quietly.
• A young, migratory raven that finds a carcass calls
loudly, attracting other migratory ravens to share the
food.
• Collective numbers of migratory birds overwhelm the
resident pair’s attempts to defend their territory.
56.6 The Evolution of Reproductive
Behavior and Mating Systems
• In sexually reproducing species, males and females
often differ in their overall reproductive strategies –
the set of behaviors that lead to reproductive success.
• Parental investment (time and energy devoted to
producing and rearing offspring) is often unequal
between the sexes.
• In the production of gametes, females contribute more
energy than males because eggs are much larger than
sperm.
Male and Female Reproductive
Strategies
• A male increases his number of offspring by mating
with multiple females, especially if he does not spend
time and energy providing parental care.
• Males may compete intensely for access to females,
and any trait that increases a male’s access or
attractiveness to females has a big reproductive payoff.
• For females, the success of her offspring may depend
on the attributes of their father or the territory he
holds.
Sexual Selection
• Male competition for access to females, and the
females’ choice of mates, establishes a form of
natural selection called sexual selection – selection
for mating success.
• Sexual selection results in males that are large, or
have ornaments and weapons (e.g., horns and
antlers) used to attract females or compete with
rival males.
• Males typically show off these elaborate structures
in complex courtship displays to attract the
attention of females.
Sexual Selection
• The degree to which females actively choose
genetically superior mates varies among species.
• In northern elephant seals, the choice is passive –
males fight violently to keep other males away.
• In sage grouse, mates are chosen by females –
males gather at display grounds (leks) where
females choose among many displaying males.
Females can be sure that offspring are theirs, but
males do not have this certainty.

Males however produce plentiful supply of sperm,

best strategy to increase their fitness = many
offspring as possible

The survival of the fittest….

Herbert Spencer 1864
Sexual Selection

• Studies of peafowl suggest a link between

ornamentation and advantageous alleles – the
offspring of fathers with impressive tails survived better
and weighed significantly more than those whose
fathers had less attractive tails.
• Another hypothesis suggests that any male that
survives despite carrying a large ornamental handicap
must have a very strong constitution indeed – and will
pass those successful alleles to his offspring.
Mating Systems

• Promiscuous species do not form close pair bonds

– both males and females mate with multiple
partners.
• In monogamous species, one male and one female
form a long-term association.
• Polygamous species have many mating partners:
• In polygyny, one male mates with many females;
• In polyandry, one female mates with multiple males.
Mating Systems

• Mating systems maximize reproductive success based

on the amount of parental care that offspring require
and other aspects of a species’ ecology.
• Species in which young require lots of parental care
from both parents (e.g., songbirds) are usually
monogamous.
• Species in which males establish large, resource-filled
territories (e.g., blackbirds) are often polygynous.
56.7 The Evolution of Social
Behavior
• Social behavior (interactions that animals have
with other members of their species) affects
reproductive success.
• Some animals are solitary, getting together only
briefly to mate (rhinoceroses and leopards), some
spend most of their lives in small family groups
(gorillas), others live in groups with thousands of
relatives (termites and honeybees).
• Some species, such as some African antelopes and
humans, live in large social units composed
primarily of nonrelatives.
Benefits of Group Living
• Groups of cooperating predators frequently capture
prey more effectively than they would on their
own.
• Prey that are subject to intense predation gain
safety in numbers, with more eyes to watch for
predators and a greater individual chance of
escaping attack.
• Some prey species, such as musk oxen, join forces
to defend themselves actively – some insects, such
as the Australian sawfly caterpillar, exhibit
cooperative defensive behavior.
Costs of Group Living
• Costs of group living include increased competition
for food, and the spread of contagious diseases and
parasites.
• Such costs are probably why the vast majority of
animals do not live in large, complex societies.
Dominance Hierarchies
• Some animal species form dominance hierarchies,
social systems in which each individual’s behavior is
governed by its place in a highly structured social
ranking.
• In a dominance hierarchy, the dominant or alpha
individual gains first access to valuable resources –
subordinate individuals concede resources to
dominant animals.
Dominance Hierarchies
• Dominant animals also incur costs – frequent
challenges from lower ranking individuals may
induce a stress response in dominant animals,
which must constantly defend their status.
• A subordinate member of a group gains benefits
such as protection against predators, and a chance
of moving up in the hierarchy as superiors age and
die.
Dominance Hierarchies

• Low-ranking males may even have a chance to

mate with one of the group’s females when
dominant males are not watching.
• Often, a subordinate’s chances of survival and
reproductive success would be lower if it lived by
itself.
Ex: Among red deer, subordinate females are at a
disadvantage in producing more prolific sons.

Primate grooming may be necessary to keep them

healthy since parasites spread easily in groups.

Humans have extensive medical care to offset health problems that

arise in living in densely populated cites
Altruism
• In some species, group members sacrifice their
own reproductive success to help individuals that
are not their direct descendants (altruism):
• Example: Subordinate members of a wolf pack do not
reproduce, but they share captured prey with the
dominant pair and that pair’s offspring.
• Altruistic behavior appears to contradict a basic
premise of Darwinian evolutionary theory – that
natural selection favors traits that increase an
individual’s relative fitness.
Degrees of Relatedness
• Helping relatives can propagate the helper’s own genes
because the family shares inherited alleles.
• The average percentage of alleles that relatives are
likely to share is determined by their degree of
relatedness.
• Full siblings share 25% of their alleles through the
mother and 25% of through the father, for a total of
50% (0.50).
• The degree of relatedness between a nephew and an
uncle is 0.25 – and between first cousins is 0.125.
Inclusive Fitness
• The costs of altruism must be measured against the
benefits of indirect reproductive success:
• Example: A young wolf reproducing on his own may
result in lower fitness (copies of alleles in the next
generation) than helping to raise his siblings.
• Inclusive fitness describes the sum of an
individual’s classical fitness (number of successful
offspring) plus fitness gained through offspring
produced by its relatives.
Kin Selection
• If altruistic behavior allows the assisted relatives to
produce proportionately more offspring than the
altruist might have produced without helping them,
the allele for altruism can increase in frequency in
the population.
• This form of natural selection is called kin
selection.
Reciprocal Altruism
• Behavioral biologists have also observed examples
of altruism between nonrelatives:
• Example: Vampire bats often share a meal with
unrelated members of their group.
• Individuals may help nonrelatives if they are likely
to return the favor in the future (reciprocal
altruism) – each member of the partnership
potentially benefits from the relationship.
Altruism in Eusocial Insects

• In some species of ants, bees, wasps, and termites

(eusocial insects), thousands of related individuals
work together in a colony for the benefit of a single
queen and her mate(s):
Example: In a honeybee colony, the only fertile female
is the queen bee – all workers (her daughters) are
sterile;
Workers perform all tasks in maintaining the hive, and
even die in defense of their colonies – why does this
self-sacrificing behavior persist over time?
Honeybees
• Female bees are diploid, receiving one set of
chromosomes from each parent.
• Male bees (drones) are haploid – they hatch from
unfertilized eggs and produce only one type of
sperm.
• Workers bees are a product of haplodiploidy, and
are related to each other by an average of 75%.
• This extremely high degree of relatedness among
workers may explain their exceptional level of
cooperation.