Chapter 8
Behavioral Ecology
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鄭先祐 生態主張者 Ayo工作室
Behavioral Ecology
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Genetic and Environmental components of Behavior
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Mechanisms of Behavioral Interaction
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Innate Behavior
Learning
Genetic and environmental factors
Communication
Aggression
Territoriality
Social systems
Environmental application
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Behavioral ecology (行為生態學)
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Behavioral ecology, the study of behavior
in an ecological context.
such as communication, aggression,
territoriality, mating systems and
cooperation.
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Genetic and environmental
components of behavior
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Innate behavior, the behaviors appear in
response to a certain stimulus and from the first
are performed in their typical stereotypical
fashion.
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The specific behavior is referred to as a fixed action
pattern (FAP).
A magnified sign stimulus that provokes an
exaggerated response is called a superstimulus.
Learning
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Ethology and comparative psychology
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The modern study of behavior has taken
two distinct routes in this century -- one
European, one American.
Fig 24.1 Two of the
bounders of modern
ethology
(a) Konrad Lorenz
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The European
approach was called
ethology. Its goal
was to understand
behavior by study its
cause, development,
evolution, and
function through the
observation of
animals in the wild,
or under somewhat
natural conditions.
Fig 24.1 Two of the
bounders of modern
ethology
(b) Niko Tinbergen
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American approach
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The American approach was called
comparative psychology.
Its goal was to understand behavior by
studying animals in the laboratory under
carefully controlled conditions.
Its focus was learning, and its primary
animal was the Norway rat.
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The definition of Instincts (本能)
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The perception of something in the
environment (a releaser) triggers a
reaction in a center in the central nervous
system(the innate releasing mechanism)
that then cause the performance of the
instinctive act, sometimes composed of
very stereotyped movements (fixed action
patterns).
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Fig. 24.2
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A male European robin in breeding
condition will attack a tuft of red feathers
(一叢紅色羽毛) placed in his territory.
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Fig. 24.4 Simplified
diagram of how a fixed
action pattern can be
triggered. The releaser
is perceived by some
sort of receptor, which
triggers the IRM to
activate certain muscles,
thereby producing an
instinctive movement
that usually involves
fixed action patterns.
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Fig. 24.3
The
scratching
movements
of dogs, as
well as
many other
vertebrates,
are
considered
fixed action
patterns.
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Learning (學習)
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Learning is a change in behavior,
based on experience.
Fig. 24.5 Chimpanzees are
highly social creatures that
live in a complex, variable,
and changing world.
Intelligence is important
under such circumstances.
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Habituation (習慣)
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Habituation is , in a sense, learning not to
respond to a stimulus.
The first time an animal encounters a
stimulus, it may respond vigorously.
But if the stimulus is presented over and
over without consequence, the response to it
gradually lessens and may finally disappear
altogether. (Fig. 24.6)
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Fig. 24.6 As animals become accustomed to a
stimulus, habituation may occur.
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Essay 24.1
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Why haven't been
selected for so that
by now we can all,
more or less,
perform such feats?
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A mnemoist
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A mnemoist (a Russian profession) would
sometimes memorize lines of 50 words.
Once, in just a few moments, he memorized
the nonsense formula
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Nd2(853/vx)(2762/n2v)(86x/273)(n2b)(rd) = sv
(1625/322)(r2s5)
Fifteen years later, upon request, he repeated
the entire formula without a single mistake.
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The interaction of genetic and
environmental factors
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The development of virtually any
behavioral trait we can imagine has both
environmental and genetic components.
譬如：Chap.2 Belding’s ground squirrel 的
kin selection. Holmes and Sherman (1982)
The behavioral interactions among
individuals in this species exemplify the
interaction between genes and the
environment.
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範例：Belding’s ground squirrel
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Holmes and Sherman (1982)
At birth, some pups were switched to different
females in combinations that created four groups of
offspring: sibs reared apart, sibs reared together,
nonsibs reared apart, and nonsibs reared together.
Whether they were sibs or not, individuals reared
together demonstrated less aggressive behavior
toward each other than individuals reared apart.
The pups apparently learn to treat the individuals
with whom they grow up as kin.
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Difference between males
and females
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Female sibs reared apart showed
lower rated of aggression toward
each other than did male sibs reared
apart.
Males were aggressive toward any
individual with whom they did not
share a nest as a juvenile.
Some innate mechanism allows
females to recognize their female kin
and behave less aggressively toward
them.
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範例：White-crowned sparrow
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Marler and Tamura (1964) showed that
males of this species have song dialects
characteristic of the region were they were
raised (Fig. 8.1)
A series of elegant experiments by Konishi
(1965) elucidated the interplay between
genetics and learning in the development
of the male song. (Fig. 8.2)
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Fig. 8.1 Song dialects in white-crowned sparrows
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Fig. 8.1 Song dialects in white-crowned sparrows
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Fig. (a) The essential events in the
development of song in male whitecrowned sparrows.
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Fig. 8.2 Two experimental manipulation of song
24 learning in male white-crowned sparrows.
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Three groups (male white-crowned sparrows)
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1. Chicks maintained in soundproof
chambers in which they were not
permitted to hear songs of any kind.(Fig.
8.2b)
2. Chicks allowed to hear songs of their
species until they were five months old, at
which point they were surgically
deafened.(Fig. 8.2c)
3. A control groups of chicks.(Fig. 8.2a)
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Mechanisms of behavioral
interaction
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Communication (溝通)
Aggression (攻擊)
Territoriality (領域)
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Communication
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Any action on the part of one organism
that alters the probability of a behavior in
another individual.
The fitness of both individuals increases
as a result.
The sending, the receiving, and the
elicited behavior are all components of the
communication system.
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化學
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聲音
視覺
接觸
四種溝通的管道
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Chemical signals
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For example, some insects use pheromones (費
洛蒙) as a sex attractants and as warning
signals (alarm pheromones).
Sex pheromones tend to be high-molecularweight compounds because these molecules
have the potential variability, and hence the
potential specificity.
Alarm pheromones tend to be volatile, lowmolecular-weight compounds which can travel
far and fast to warn conspecifics..
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Discrete and graded signals
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Communication signals can be divided into two
basic types: discrete signals and graded signals.
Discrete signals provide simple, digital– yes
or no, on or off – kinds of information.
Graded signals fall along a continuum of
intensity or complexity, and their variability
allows the signaler to provide richer, analog
information.
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範例：螢火蟲(Fig. 3)
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Produce discrete signals.
Each species has a characteristic flashing
pattern that is relatively invariant(Fig. 8.4).
The flashing patterns of both males and females
are highly stereotyped.
Males fly across the habitat and flash.
The female communicates her receptivity and
location by responding with her own flashing.
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Fig. 8.4 Discrete
signals in the
sexual
communication of
fireflies.
When a female on
the ground
observes the
pattern of her own
species, she
flashes in
response,
attracting the male
to her.
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範例：
the waggle dance of the 蜜蜂
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A series of graded communication signals.
When a honeybee locates a new food source,
she returns to the hive and communicates the
direction, distance, and quantity of food to her
nest mates.
In performing the waggle dance, depicted in Fig.
8.5, the worker bee repeatedly moves in a
figure-eight, she performs a so-called straight
run, a straight-line movement during which she
vibrates(waggles) her abdomen.
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Fig. 8.5 The waggle dance of the honeybee, which is
performed both (a) outside the hive and (b) inside the
hive on a vertical surface.
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Fig. 8.5 The waggle dance of the honeybee, which is
performed both (a) outside the hive and (b) inside the
hive on a vertical surface.
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There is a
significant
positive
relationship
between the
complexity of
the social
system and the
size of the
alarm call
repertoire.
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Aggression
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Aggression, as an act by which one
individual achieves dominance over another
by physical violence or the threat of it.
Such behavior is typically considered an
aspect of predation, defense against
predators, or competition.
不戰而屈人之兵，謂之上策。
Fighting is more ceremonial than lethal.
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儀式化的攻擊
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Ritualized pattern evolved because the
fitness of the species depends on
protecting important members from lethal
combat that would deprive them of the
opportunity to pass on their genes.
或許可以用 Group selection argument.
但是 the advantage for the group is
different from what is advantageous for the
individual.
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增進個人的fitness
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It is more likely that the adaptive basis for
ritualization of such behavior involves individual
fitness.
It may be to both antagonists’ advantage to
recognize quickly, and with minimum trauma.
The younger or smaller individual often loses
such an encounter, but given time, that
individual may well become large enough to
dominate others.
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範例：the great tit (大山雀)和
紅翅黑鳥(red-winged blackbirds)
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大山雀可以發出多達八種宣告領域的叫聲，
這會讓沒有領域者誤以為有許多佔有領域
者在這個地區，而離開該地區。
相似的結果也發生在紅翅黑鳥，運用錄音
播放多隻有領域者的叫聲，入侵此地區者
就會較少。若是只有單隻的叫聲，入侵者
就會較多。
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範例：the spotted hyenas (土狼)
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One of the most violently aggressive mammals.
Intense competition and aggression occur over
kills that the pack has made.
In response to selective pressure, females are
larger and more aggressive than males,
presumably to ensure access to kills for
themselves and their offspring.
Females also are anatomically masculinized(雄
性化), which was achieved via increased
androgen levels.
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範例：infanticide (殺嬰)
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It appears that in the males of the whitefooted mouse, infanticide is associated
with confidence of paternity just as it it in
lions.
In females, infanticide appears to be
related to crowding.
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Territoriality (領域現象)
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Home range (Fig. 8.8)
Territory (領域)：Territorial behavior
evolves to ensure access to crucial
resources.
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3D的Home
Range
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Classification of territories
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Type A：Large defended area; courtship, mating,
nesting, and food gathering.
Type B：Large defended area; all breeding but
not foraging.
Type C：Small defended area around nest.
Type D：Mating territories; for courtship
activities only.
Type E：Roosting or shelter positions.
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The size of a territory
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領域雖然愈大可能愈好，但是要維持愈大
的領域，所要花費的投資將會愈大。
擁有領域者，受到被掠食的風險也加大。
Type A的物種，其size受到food supply的
量影響。食物供應愈缺乏，松鼠的領域通
常會愈大。(Tamiasciurus squirrels)
Energy budget of territorial African goldenwinged sunbirds (Table 8.3)
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If the production rate
of a defended territory
is twice that of an
undefended site, the
bird gains 2,400K per
day by maintaining
the territory .
休息與覓食，兩
者能量的耗用相
差600K/小時。
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Territory size
increases as a
function of body
mass.
The slope is
greater for
carnivores.
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Social systems
Three fundamental components:
 1. Group size and structure
 2. The mating system
 3. Cooperation and helping
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Group size and structure
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An aggregation is a group of conspecifics
that includes more than one family unit
and is localized in a particular area.
A colony is a group of conspecifics that
form a society of highly integrated
individuals.
Group. This term has two meanings. 一個
是泛稱。另一個是指介乎在aggregation和
colony之間。
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The evolutionary “decision”
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The evolutionary “decision” about whether
to join a group is based on the relative
advantages of solitary living and group
living.
An example of the balance that must be
struck in the evolution of group living – the
optimization of group size – is shown in
Figure 8.10.
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The optimal
group size is
found at the
intersection of
those two
functions.
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“A” might be
defense against
predators.
小團體，個體少
不足以防衛。
過大的團體，招
惹掠食者的攻擊。
“B” might
represent group
foraging.
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Mating system
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The relationships among males and
females during breeding as the mating
system.
Three components:
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1. The number of mates an individual takes.
2. Whether the male and female form a pair
bond.
3. The duration of the pair bond.
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Polygyny
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Males mate with more than one female.
This implies that not all males mate, there is
variance in male reproductive success.
If resources, leading to intense competition
among the males. Because females choose
males who control resources, there is a high
variance in reproductive success among males,
and the potential for polygyny arises. (Emlen
and Oring’s model)
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Synchrony in sexual receptivity
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As the degree of synchrony in sexual receptivity
of females decreases, the opportunity for
multiple matings increases.
If all females are receptive at the same time, it
will be difficult for a male to service more than
one.
At high levels of asynchrony, the cost of
maintaining exclusive use of a resource exceeds
the potential reproductive benefits. 因此
polygyny decreases again. (Fig. 8.12)
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Polygyny 的產生，
受到兩項因素的
影響：
1. Asynchrony 的
程度
2. 資源clumped
的程度。
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Female defense polygyny
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When females associate with each other
in groups.
Most often the selective pressure for
female associations is predation.
例如：In gorillas, females travel in groups
to avoid leopard predation; males defend
such groups from other males.
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Resource defense polygyny
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Resource is clumped, and males defend the
resource.
Females are attracted to the territory for its
resource, and males gain access to multiple
females in this way.
例如：the orange-rumped honeyguid, a bird of
Africa and the Indian subcontinent, feeds
entirely on beeswax. Males defend bee
colonies from other males. Females are
attracted to the colonies and copulate with the
male. Extreme polygyny results.
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Lek systems
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In lek systems, the males defend only a tiny portion
of a communal display area.
Females approach the display area and choose
among the males that are displaying there.
例如：sage grouse leks in prairie and sage habitats,
as many as 400 males may display over an area of
more than a hectare. Fewer than 10% of the males
achieve 75% of the matings.
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Monogamy
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The frequency of monogamy is far higher in
birds than in mammals; approximately 90% of all
bird species are monogamous. In contrast,
fewer than 5% of mammals are monogamous.
雄性多次 matings 的 fitness advantage 非常的大。
因此必要有狀況的限制，才能避免其polygyny。
鳥類與哺乳類的monogamy發生率的差異，顯現
出限制狀況的差異。
鳥類因為the timing of resource availability的限制，
雌性傾向於生殖synchrony。
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Polyandry
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Polyandry, in which females mate with more
than one male, is the rarest of the mating
systems.
In some cases of polyandry, the sex roles are
reversed, and males provide most of the
parental care.
譬如：The American jacana(Jacana spinosa), a
wading bird, exhibits an extreme form of
polyandry. Females defend large “superterritories” that contain multiple nests attended
by different males.
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The spotted sandpiper (Actitis macularia)
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Females compete for mating territories,
into which two or three males settle.
Each male defends a single nest in a
portion of the female’s territory, providing
parental care for the young, while the
female defends her territory and attempts
to attract still other mates.
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Cooperation and helping
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Altruism (利他行為)
Genetically based cooperation
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Habitat saturation and food supply
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Kin relationships.
Coefficient of relationship
A number of example of cooperative breeding
have evolved among the birds.
One of the most thoroughly studies is the
acorn woodpecker.
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The acorn woodpecker
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The social unit of this species is a
territorial group of up to a dozen
members(both males and females) that
gather and store acorns in thousands of
holes drilled in a tree called a granary.
This cache provides the major source of
food over the winter.
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Three unusual features(Fig. 8.14)
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1. Division of labor and helpers
2. Mate sharing
3. Infanticide. Females kill the first eggs
laid by other females. Whenever birds
enter a group to fill a vacancy, they kill all
the young.
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圖8.14 The structure of the acorn woodpecker
group is strongly influenced by the quantity of
mast the group can store in the fall.
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Why?
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1. The habitat is saturated; all good territories
are occupied. Consequently, opportunities for
young to disperse are extremely limited.
2. Mate sharing results because it provides
advantages to the individual breeder.
3. The infanticide is apparently adaptive
because it provides a genetic advantage over
other members of the group.
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Eusocial system
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1. Extensive cooperation
2. Division of labor
3. The presence of a queen.
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Fig. 8.17 Relationships between (a) reproductive costs and (b)
reproductive benefits of breeders in relation to the amount of
parental care.
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Environmental Application
Behavioral Ecology and captive breeding
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運用captive breeding 繁殖，
以增長滅絕中的生物族群。
運用過此方法的有：
California condor, blackfooted ferret and the golden
lion tamarin of the Brazilian
rain forests.
成功的條件：了解其行為和
相關社會因素。
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例如：Wolf packs and lion prides
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The groups of both species are similar in
size and composition.
Wolves are basically monogamous, and
only the alpha male and alpha female
mate. Other adults participate in rearing
the offspring but do not breed.
Lion prides contain several breeding
females. The males are usually related to
each other.
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 問題與討論！
ayo@faculty.pccu.edu.tw
Ayo 文化站 http://faculty.pccu.edu.tw/~ayo
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