Chapt 32

669
Ecology began as a descriptive

science that has now become an
experimental and predictive sci-
ence. Models have been devel-
oped that predict how population
sizes within communities change
over time due to species inter-
actions. Some ecologists study
and also include in their models
physical factors that inuence
energy ow and nutrient cycling
within an ecosystem. They know
that humans alter the transfer
rates of substances within biogeo-
chemical cycles that maintain the
biosphere. This accounts for acid
rain, global warming, ozone
depletion, and other changes that
are expected to adversely affect all
species.
Behavior and Ecology
32 Animal Behavior 671
Animals respond to stimuli in keeping
with their own neurophysiology and in a
manner that increases the likelihood of
their survival and ability to reproduce.
34 Community Ecology 701
Species in a particular locale make up a
community where interactions include
competition, predation, and various
symbiotic relationships.
36 Ecosystems and Human
Interferences 743
An ecosystem, which consists of a
community and its physical environment,
is characterized by energy ow and
chemical cycling.
33 Population Ecology 687
The distribution and abundance of a
species may be related to its life history
pattern in order to predict its future
population size.
35 Biosphere 717
Terrestrial and aquatic communities are
located within the biosphere, that portion
of the earths crust, atmosphere, and
waters, where living things exist.
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Ethologists study the behavior of animals in the wild. It is one of
the most difcult and rewarding of elds, because it requires the
study of animals in their own territory over extended periods of
time. Such study requires patience, stamina, and creativity. Etholo-
gists may study behaviors such as mating, territorial protection,
feeding, migration, and hibernation.
Ecologists study the relationships between living things and their
environment, and among different species. They measure the
importance of such interactions to the species being studied. The
results of these studies help humans to understand the signicance
of development on natural areas.
Environmental engineers are multidisciplinary specialists who
help preserve the environment and plan the cleanup of polluted
areas. These engineers use experience in engineering, chemistry,
and biology to determine the extent of the pollution and the costs
and benets of different levels of cleanup effort. They also advise
developers regarding the potential impact of new projects on the
environment. Many also work to devise better ways to limit air and
water pollution by industry.
Wildlife managers determine which species should be available
for hunters and shermen to continue their sports. They identify
the behavioral and environmental factors that affect population
sizes and develop management programs that will permit a sus-
tained yield. Endangered species management also falls under
wildlife management, but in this case, the species are protected and
are not to be killed by anyone.
Foresters manage, develop, and help protect forest resources.
Foresters manage timberland, which involves a variety of duties.
Those working in private industry may be responsible for procur-
ing timber from private landowners. Foresters also supervise the
planting and growing of new trees. They advise on the type, num-
ber, and placement of trees to be planted. Foresters monitor the
trees to ensure healthy growth and to determine the best time for
harvesting. If foresters detect signs of disease or harmful insects,
they decide on the best course of treatment to prevent contamina-
tion or infestation of healthy trees.
Behaviorist collaring a coyote.
Ecologists measuring tropical trees.
Environmental engineer testing for harmful gases.
Careers in Ecology
670
Animal Behavior
Chapter Concepts
An animal is organized to carry out behaviors
that help it survive and reproduce. 672
32.1 Genetic Basis of Behavior
Behaviors have a genetic basis but can also be
inuenced by environmental factors. 672
The nervous and endocrine systems have
immediate control over behaviors. 673
32.2 Development of Behavior
Behaviors sometimes undergo development after
birth, as when learning affects behavior. 674
32.3 Adaptiveness of Behavior
Natural selection inuences such behaviors as
methods of feeding, selecting a home, and
reproducing. 676
32.4 Animal Societies
Animals living in societies have various means
of communicating with one another. 680
32.5 Sociobiology and Animal Behavior
Apparently, altruistic behavior only occurs if it
actually benets the animal. 683
These western grey kangaroos, Macropus fulginosus, are trying
to push each other to the ground in a battle over females. Aggres-
sion between members of a society is ritualized and neither party
in the struggle is usually harmed by the conict.
671
The behavior of animals is any action that can be observed
and described. All behavior has a genetic base in that the
anatomy and physiology of the animal which is inherited is
suitable to performing the behavior. In addition, various ex-
periments have been done to show that specic behaviors
have a genetic base.
A peach-faced lovebird, Agapornis roseicollis, cuts long,
regular strips of material with its strong beak and then tucks
them in its rump feathers for transport to the nest. AFishers
lovebird, Agapornis scheri, carries stronger materials, such
as sticks, directly in its beak.
672 Part 7 Behavior and Ecology 32-2
A
t the start of the breeding season, male bowerbirds
use small sticks and twigs to build elaborate display
areas called bowers. They clear the space around the
bowers, removing leaves and debris, and decorate the area
with fresh owers, fruits, moss, mushrooms, pebbles, or
shells. Each species has its own preference in decorations.
The satin bowerbird of eastern Australia prefers blue ob-
jects, a color that harmonizes with the males glossy blue-
black plumage (Fig. 32.1).
After the bower is complete, a male spends most of his
time near his bower, calling to females, renewing his decora-
tions and guarding his work against possible raids by other
males. After inspecting many bowers and their owners, a fe-
male approaches one and the male begins a display. He
faces her, uffs up his feathers, and aps his wings to the
beat of a call. The female enters the bower, and if she
crouches, the two mate.
Ethologists (scientists who study behavior) want to
know how the male bowerbird is structured to perform this
behavior and how the behavior helps him secure a mate.
In general, ethologists determine how a behavior is con-
trolled and how a behavior enables an animal to survive
and/or reproduce.
If the behavior for obtaining and carrying nesting mate-
rial is inherited, then hybrids might show intermediate be-
havior. When the two species of birds were mated, it was
observed that the hybrid birds have difculty carrying nest-
ing materials. They cut strips and try to tuck them in their
rump feathers, but they are unsuccessful. After a long period
of time (about three years), a hybrid learns to carry the cut
strips in its beak but still briey turns its head toward its
rump before ying off. Therefore, these studies support the
hypothesis that behavior has a genetic basis.
Several experiments have been done with the garter
snake, Thamnophis elegans, which has two different types of
snake populations in California. Inland populations are
aquatic and commonly feed underwater on frogs and sh.
Coastal populations are terrestrial and feed mainly on slugs.
In the laboratory, inland adult snakes refused to eat slugs
while coastal snakes readily did so. To test for possible ge-
netic differences between the two populations, matings
were arranged between inland and coastal individuals, and
it was found that isolated newborns show an overall inter-
mediate incidence of slug acceptance.
The difference between slug acceptors and slug rejecters
appears to be inherited, but what physiological difference
have the genes brought about? A clever experiment an-
swered this question. When snakes eat, their tongues carry
chemicals to an odor receptor in the roof of the mouth. They
Figure 32.1 Mating behavior of satin bowerbirds.
A female satin bowerbird has chosen to mate with this male. Most
likely, she was attracted by his physique and the blue decorations of
his bower.
A. roseicollis A. fischeri
use tongue icks to recognize their prey. Even newborns will
ick their tongues at cotton swabs dipped in uids of their
prey. Swabs were dipped in slug extract, and the number of
tongue icks were counted for newborn inland and coastal
snakes. Coastal snakes had a higher number of tongue icks
than inland snakes (Fig. 32.2). Apparently, inland snakes do
not eat slugs because they are not sensitive to their smell. A
genetic difference between the two populations of snakes
has resulted in a physiological difference in their nervous
systems. Although hybrids showed a great deal of variation
in the number of tongue icks, they were generally interme-
diate as predicted by the genetic hypothesis.
Both nervous and endocrine systems are responsible
for the integration of body systems. Is the endocrine sys-
tem also involved in behavior? Various studies have been
done to show that it is. For example, the egg-laying behav-
ior in the marine snail Aplysia involves a set sequence of
movements. Following copulation, the animal extrudes
long strings of more than a million egg cases. It takes the
egg case string in its mouth, covers it with mucus, waves
its head back and forth to wind the string into an irregular
mass, and attaches the mass to a solid object, like a rock.
Several years ago, scientists isolated and analyzed an egg-
laying hormone (ELH) that causes the snail to lay eggs
even if it has not mated. ELH was found to be a small pro-
tein of 36 amino acids that diffuses into the circulatory sys-
tem and excites the smooth muscle cells of the
reproductive duct, causing them to contract and expel the
egg string. Using recombinant DNA (deoxyribonucleic
acid) techniques, the investigators isolated the ELH gene.
The genes product turned out to be a protein with 271
amino acids. The protein can be cleaved into as many as 11
possible products, and ELH is one of these. ELH alone, or
in conjunction with these other products, is thought to con-
trol all the components of egg-laying behavior in Aplysia.
The results of many types of studies support the
hypothesis that behavior has a genetic basis and
that genes inuence the development of neural
and hormonal mechanisms that control behavior.
Chapter 32 Animal Behavior 673 32-3
Inland garter snake does
not eat slugs.
Coastal garter snake does
eat slugs.
20
15
inland
10
1.0 2.0 3.0 4.0 5.0
5
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Tongue Flicks per Minute
coastal
Figure 32.2 Feeding behavior of garter snakes.
The number of tongue icks by inland and coastal garter snakes as a response to slug extract on cotton swabs. Coastal snakes tongue-icked
more than inland snakes.
Given that all behaviors have a genetic basis, we can go on
to ask if environmental experiences after hatching or birth
also shape the behavior. Some behaviors seem to be stereo-
typedthey are always performed the same way each time.
These were called xed action patterns (FAP), and it is said
that FAPs were elicited by a sign stimulus, a cue that sets the
behavior in motion. For example, human babies will smile
when a at but face-sized mask with two dark spots for eyes
is brought near them. Its possible that some behaviors are
FAPs, but increasingly, investigators are nding that many
behaviors, formerly thought to be FAPs, develop after
practice.
Laughing gull chicks begging behavior is always per-
formed the same way in response to the parents red beak. A
chick directs a pecking motion toward the parents beak,
grasps it, and strokes it downward (Fig. 32.3). Sometimes a
parent stimulates the begging behavior by swinging its beak
gently from side to side. After the chick responds, the parent
regurgitates food onto the oor of the nest. If need be, the
parent then encourages the chick to eat. This interaction be-
tween the chicks and their parents suggests that the begging
behavior involves learning. (Learning is dened as a
durable change in behavior brought about by experience.)
To test this hypothesis, diagrammatic pictures of gull heads
were painted on small cards and then eggs were collected in
the eld. The eggs were hatched in a dark incubator to elim-
inate visual stimuli before the test. On the day of hatching,
each chick was allowed to make about a dozen pecks at the
model. The chicks were returned to the nest and were each
retested. The tests showed that on the average, only one-
third of the pecks by a newly hatched chick strike the model.
But one day after hatching, more than half of the pecks are
accurate, and two days after hatching, the accuracy reaches
a level of more than 75%. Investigators concluded that im-
provement in motor skills, as well as visual experience,
strongly affect development of chick begging behavior.
Behavior has a genetic basis, but the development
of mechanisms that control behavior is subject to
environmental inuences, such as practice after
birth.
How do chicks recognize a parent? Newly hatched
chicks peck equally at any model as long as it has a red beak.
Chicks a week old, however, will peck only at models that
closely resemble the parent. Perhaps operant conditioning
with a reward of food could account for this change in be-
havior. Operant conditioning, which is one of many forms
of learning, is often dened as the gradual strengthening of
stimulus-response connections. In everyday life, most peo-
ple know that animals can be taught tricks by giving re-
wards such as food or affection. The trainer presents the
stimulus, say a hoop, and then gives a reward (food) for the
proper response (jumping through the hoop). B. F. Skinner is
well known for studying this type of learning in the
Pecking
accuracy
of newborn
Pecking
accuracy of
two-day old
Mean accuracy of pecking model
for all chicks tested
Days in nest
H
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(
p
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)
0
25
50
75
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Figure 32.3 Pecking behavior of laughing gulls.
At about three days, a laughing gull chick grasps the beak of a
parent, stroking it downward, and the parent then regurgitates food.
laboratory. In the simplest type of experiment performed by
Skinner, a caged rat happens to press a lever and is re-
warded with sugar pellets, which it avidly consumes. There-
after, the rat regularly presses the lever whenever it is
hungry. In more sophisticated experiments, Skinner even
taught pigeons to play ping-pong by reinforcing desired re-
sponses to stimuli.
Imprinting is another form of learning; chicks, ducklings,
and goslings will follow the rst moving object they see after
hatching. This object is ordinarily their mother, but they
seemingly can be imprinted on any objecta human or a red
ballif it is the rst moving object they see during a sensitive
period of two to three days after hatching. The term sensitive
period means that the behavior only develops during this
time. Although the Englishman Douglas Spalding rst ob-
served imprinting, the Austrian Konrad Lorenz is well
known for investigating it. He found that imprinting not only
served the useful purpose of keeping chicks near their
mother, it also caused male birds to court a member of the cor-
rect speciessomeone who looks like mother! The goslings
who had been imprinted on Lorenz courted human beings
later in life. In-depth studies on imprinting have shown that
the process is more complicated than originally thought. Eck-
hard Hess found that mallard ducklings imprinted on hu-
mans in the laboratory would switch to a female mallard that
had hatched a clutch of ducklings several hours before. He
found that vocalization before and after hatching was an im-
portant element in the imprinting process. Female mallards
cluck during the entire time that imprinting is occurring. Do
social interactions inuence other forms of learning? Patterns
of song learning in birds suggests that they can.
Song Learning in Birds
During the past several decades, an increasing number of
investigators have studied song learning in birds. White-
crowned sparrows sing a species-specic song, but males
of a particular region have their own dialect. Birds were
caged in order to test the hypothesis that young white-
crowned sparrows learn how to sing from older members
of their species (Fig. 32.4). A group of birds that heard no
songs at all sang a song, but it was not fully developed.
Birds that heard tapes of white-crowns singing sang in that di-
alect, as long as the tapes were played during a sensitive
period from about age 10 to 50 days. White-crowned spar-
rows dialects (or other species songs) played before or af-
ter this sensitive period had no effect on their song.
Apparently, their brain is especially primed to respond to
acoustical stimuli during the sensitive period. Neurons
that are critical for song production have been located, and
they re when the birds own song is played or when a
song of the same dialect is played. Other investigators have
shown that birds given an adult tutor will sing the song of
even a different speciesno matter when the tutoring be-
gins! It would appear that social experience has a very
strong inuence over the development of singing.
Animals have an ability to benet from experience;
learning occurs when a behavior changes with
practice.
Isolated bird sings
but song is not developed.
Bird sings developed song played
during a sensitive period.
Bird sings song of social tutor without regard to
sensitive period.
Figure 32.4 Song learning by white-crowned sparrows.
Three different experimental procedures are depicted and the results noted. These results suggest that there is both a genetic basis and an
environmental basis for song learning in white-crowned sparrows.
Since genes inuence the development of behavior, it is rea-
sonable to assume that behavioral traits (like other traits) are
subject to natural selection. Our discussion will focus on re-
productive behaviorspecically, the manner in which ani-
mals secure a mate. But we will also touch on the other two
survival issuescapturing resources and avoiding preda-
torsbecause these help an animal survive, and without
survival, reproduction is impossible. Investigators studying
survival value seek to test hypotheses that specify how a
given trait might improve reproductive success.
Males can father many offspring because they continu-
ally produce sperm in great quantity. We would then expect
competition among males to inseminate as many different
females as possible. In contrast, females produce few eggs,
so the choice of a mate becomes a prevailing consideration.
Sexual selection can bring about evolutionary changes in the
species. Sexual selection is changes in males and females,
often due to male competition and female selectivity, lead-
ing to reproductive success.
Female Choice
Courtship displays are rituals that serve to prepare the sexes
for mating. They help male and female recognize each other
so that mating will be successful. They also play a role in a
females choice of a mate.
In a study of satin bowerbirds
(see Fig. 32.1), two oppos-
ing hypotheses regarding
female choice were tested:
Good genes hypothesis: females
choose mates on the basis
of traits that improve their
chances of survival.
Run-away hypothesis: females
choose mates on the basis
of traits that make them
attractive to females. The
term run away pertains
to the possibility that the
trait will be exaggerated in
the male until its reproduc-
tively favorable benet is
checked by the traits
unfavorable survival cost.
Investigators watched bowerbirds
at feeding stations and also moni-
tored the bowers. They discovered
that although males tend to steal
blue feathers and/or actively de-
stroy a neighbors bower, more ag-
gressive and vigorous males were
able to keep their bowers in good condition. These were the
males usually chosen as mates by females. These data do not
clearly support either hypothesis. It could be that aggres-
siveness, if inherited, does improve the chances of survival,
or it could be that females simply preferred bowers with the
most blue feathers.
The raggiana bird of paradise is remarkably
dimorphicthe males are larger than females and have
beautiful orange ank plumes. In contrast, the females are
drab (Fig. 32.5). Courting males, which form a group called
a lek, gather and begin to call. If a female joins them, the
males raise their orange display plumes, shake their wings
and hop from side to side, while continuing to call. They
then stop calling and lean upside down with the wings pro-
jected forward to show off their beautiful feathers.
Female choice can explain why male birds are so much
more showy than females. The remarkable plumes of the
male might signify health and vigor to the female, just as a
well-constructed bower might. In barn swallows, females
also choose those with the longest tails and investigators
have shown that males relatively free of parasites have
longer tails than otherwise.
Is a difference in reproductive behavior related to a par-
ticular food source? Raggiana birds forage far and wide for
their food (nutritious, complex fruits), and lekking is one
way for males to attract the wide-ranging females. The male
raggiana is polygynous (has more than one mate) and does
not help raise the offspring. On the other hand, a related
male
fruit
mahogany tree
female
Figure 32.5 Mating behavior in birds of paradise.
In birds of paradise, males have resplendent plumage brought about by sexual selection. The
females are widely scattered, foraging for complex fruits; the males form leks that females visit to
choose a mate.
species, the trumpet manucode, Manucodia keraudrenit, feeds
on gs, which are more prevalent but not as nutritious as
complex fruits. These birds are monogamousthe pair
bonds perhaps for life, and the males are not as colorful as
raggiana males. Both sexes are needed to successfully raise
the young, and this relaxes pressure on the male to be showy.
Male Competition
Studies have been done to determine if the benet of mating
is worth the cost of competition among males. Only if the
positive effects outweigh the negative effects will the animal
enjoy reproductive success.
Dominance Hierarchy
Baboons, a type of Old World monkey, live together in a
troop. Males and females have separate dominance hierar-
chies in which a higher ranking animal has greater access to
resources than a lower ranking animal. Dominance is de-
cided by confrontations, resulting in one animal giving way
to the other.
Baboons are dimorphic; the males are larger than the fe-
males, and they can threaten other members of the troop
with their long, sharp canines (Fig. 32.6). The baboons travel
within a home range, foraging for food each day and sleep-
ing in trees at night. The dominant males decide where and
when the troop will move and, if the troop is threatened,
they cover the troop as it retreats and attack intruders when
necessary.
Females undergo a period known as estrus, during
which they ovulate and are willing to mate. At this time a fe-
male approaches a dominant male and they form a mating
pair for several hours or days. The male baboon pays a cost
for his dominant position. Being larger means that he needs
more food, and being willing and able to ght predators
means that he may get hurt, and so forth. Is there a repro-
ductive benet to his behavior? Yes, in that dominant males
do indeed monopolize estrous females when they are most
fertile (Figure 32.7).
Nevertheless, there are other avenues to fathering off-
spring. Some males act as helpers to particular females and
her offspring; the next time she is in estrus she may mate
preferentially with him instead of a dominant male. Or sub-
ordinate males may form a friendship group that can op-
pose a dominant male, making him give up a receptive
female.
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Rank of Male in Dominance Hierarchy
1 2 3 4 5 6 7 8 9 10
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Figure 32.6 A male olive baboon displaying full threat.
In olive baboons, males are larger than females and have enlarged
canines. Competition between males establishes a dominance
hierarchy for the distribution of resources.
Figure 32.7 Female choice and male dominance among
baboons.
Although it may appear that females mate indiscriminately, they mate
more often with a dominant male when they are most fertile.
Miniature radio transmitters that emit a radio signal allow you
to track an animal in the wild. The transmitter is encapsulated
along with a battery in a protective epoxy resin covering, and
the package is either attached to an animal with a collar or a clip,
or implanted surgically into the main body cavity of the animal.
In either case, it is important to capture the animal carefully to
use sedation or anesthesia to calm the animal during tagging or
surgical implantation. The animal is permitted to recover fully
prior to its release. A radio transmitter device is generally no
more than 10% of the animals body weight, and therefore it
should not interfere with normal activities.
To track an animal, a researcher needs a radio receiver
equipped with an antenna and earphones. The strongest signal
comes from the direction of the animal as the antenna is rotated
above the head. The data obtained allow the researcher to obtain
a series of xes (to determine where the animal is) and calculate
its rate of movement. Plotting the sequence of xes on a map of
the appropriate scale gives information on the area that is used
by the animal in the course of a night or several nights (Fig. 32A).
The distance over which the signal travels varies with the
size of the transmitter and the strength of the battery. For small
rodents it may be necessary to be within 510 meters of the ani-
mal to hear the signal; for larger collars like some of those used
on elk or caribou, the signal can actually be received by a satel-
lite orbiting above the earth. Later, the signals are sent in a radio
beam by antenna to the ground. Satellite transmission allows re-
searchers to work with animals that are far away, and prevents
any possibility of interfering with the animals activities.
Anumber of important ndings have been made utilizing ra-
diotelemetry, and a few examples provide a avor of the kinds
of results obtained with this technology. Elk in Yellowstone Na-
tional Park in Wyoming have summer home range areas that are
510 times larger than in winter. Areas occupied by many mice
and voles get larger as the animal ages, from the time the ani-
mals are weaned until they are adults, but these increases are
greater for males than for females. Winter dens of rattlesnakes
(and other snakes) are often some distance from the areas where
they spend their summer months. Some crows migrate each
spring and fall, whereas others remain as residents in the same
locale throughout the year.
Radio transmitters can also be used to obtain data on physio-
logical functions (heart rate, body temperature) in conjunction
with observations of behavior while animals continue to engage
in normal activities. When physiological measurements are
taken, a device is included in the package that is sensitive to tem-
perature or heart rate and can translate that information to the
transmitter. Altogether, the use of radio transmitters has added a
new dimension to the ability to explore the lives of animals.
678
Tracking an Animal in the Wild
b. Plot of activity range
a. Radiotransmitter system
antenna
signal
collar with
transmitter
receiver
10 m
Figure 32A Electronic animal tracking.
a. By using a miniature radio transmitter in a neck collar, an antenna, and a receiver, it is possible to record an animals location
approximately every 15 minutes throughout a 24-hour period. b. These data are then used to plot its activity range for that day. (Some data
in the plot represent multiple xes at the same location.)
Territoriality
A territory is an area that is defended against competitors.
Territoriality includes the type of defensive behavior
needed to defend a territory. Vocalization and displays,
rather than outright ghting, may be sufcient to defend a
territory (Fig. 32.8). Male songbirds use singing to announce
their willingness to defend a territoryso other males of the
species become reluctant to make use of the same area.
Red deer stags (males) on the Scottish island of Rhum
compete to be the harem master of a group of hinds (fe-
males) that mate only with them. The reproductive group
occupies a territory that the harem master defends against
other stags. Harem masters rst attempt to repel challengers
by roaring. If the challenger remains, the two lock antlers
and push against one another. If the challenger now with-
draws, the master pursues him for a short distance, roaring
the whole time. If the challenger wins, he becomes the
harem master.
Aharem master can father two dozen offspring at most,
because he is at the peak of his ghting ability for only a
short time. And there is a cost to be able to father offspring.
Stags must be large and powerful in order to ght; therefore,
they grow faster and have less body fat. During bad times,
they are more likely to die of starvation, and in general, they
have shorter lives. The behavior of harem defense by stags
will only persist in the population if its cost (reduction in the
potential number of offspring because of a shorter life) is
less than its benet (increased number of offspring due to
harem access).
Evolution by sexual selection can occur when
females have the opportunity to select among
potential mates and/or when males compete
among themselves for access to reproductive
females.
Figure 32.8 Competition between males among red deer.
Male red deer compete for a harem within a particular territory. a. Roaring alone may frighten off a challenger. b. But outright ghting may be
necessary, and the victor is most likely the stronger of the two animals.
a.
b.
There is a wide diversity of social behavior among animals.
Some animals are largely solitary and join with a member of
the opposite sex only for the purpose of reproduction. Others
pair, bond, and cooperate in raising offspring. Still others form
a society in which members of species are organized in a coop-
erative manner, extending beyond sexual and parental behav-
ior. We have already had occasion to mention the social groups
of baboons and red deer. Social behavior in these and other
animals requires that they communicate with one another.
Communicative Behavior
Communication is an action by a sender that inuences the
behavior of a receiver. The communication can be purposeful
but does not have to be purposeful. Bats send out a series of
sound pulses and listen for the corresponding echoes in order
to nd their way through dark caves and locate food at night.
Some moths have an ability to hear these sound pulses, and
they begin evasive tactics when they sense that a bat is near.
Are the bats purposefully communicating with the moths? No,
bat sounds are simply a cue to the moths that danger is near.
Communication is an action by a sender that
affects the behavior of a receiver.
Chemical Communication
Chemical signals have the advantage of working both night
and day. The term pheromone is used to designate chemical
signals in low concentration that are passed between mem-
bers of the same species. Female moths secrete chemicals
from special abdominal glands, which are detected down-
wind by receptors on male antennae. The antennae are espe-
cially sensitive, and this assures that only male moths of the
correct species (and not predators) will be able to detect them.
Cheetahs and other cats mark their territories by
depositing urine, feces, and anal gland secretions at the
boundaries (Fig. 32.9). Klipspringers (small antelope) use se-
cretions from a gland below the eye to mark twigs and
grasses of their territory.
Auditory Communication
Auditory (sound) communication has some advantages
over other kinds of communication (Fig. 32.10). It is faster
than chemical communication, and it also is effective both
night and day. Further, auditory communication can be
modied not only by loudness but also by pattern, duration,
and repetition. In an experiment with rats, a researcher dis-
covered that an intruder can avoid attack by increasing the
frequency with which it makes an appeasement sound.
Male crickets have calls, and male birds have songs for a
number of different occasions. For example, birds may have
one song for distress, another for courting, and still another
for marking territories. Sailors have long heard the songs of
humpback whales because they are transmitted through the
hull of a ship. But only recently has it been shown that the
song has six basic themes, each with its own phrases, that
can vary in length and be interspersed with sundry cries and
chirps. The purpose of the song is probably sexual and
serves to advertise the availability of the singer. Language is
the ultimate auditory communication, but only humans
Figure 32.9 Use of pheromone to mark a territory.
This male cheetah is spraying a pheromone onto a tree in order to
mark his territory.
Figure 32.10 A chimpanzee with a researcher.
Chimpanzees are unable to speak but can learn to use a visual
language consisting of symbols. Some believe chimps only mimic
their teachers and never understand the cognitive use of a language.
Here the experimenter shows Nim the sign for drink. Nim copies.
have the biological ability to produce a large number of dif-
ferent sounds and to put them together in many different
ways. Nonhuman primates have at most only forty different
vocalizations, each having a denite meaning, such as the
one meaning baby on the ground, which is uttered by a
baboon when a baby baboon falls out of a tree. Although
chimpanzees can be taught to use an articial language, they
never progress beyond the capability level of a two-year-old
child. It has also been difcult to prove that chimps under-
stand the concept of grammar or can use their language to
reason. It still seems as if humans possess a communication
ability unparalleled by other animals.
Visual Communication
Visual signals are most often used by species that are active
during the day. Contests between males make use of threat
postures and possibly prevent outright ghting that might
result in reduced tness. A male baboon displaying full
threat is an awesome sight that establishes his dominance
and keeps peace within the baboon troop (see Fig. 32.6).
Hippopotamuses have territorial displays that include
mouth opening.
The plumage of a male raggiana bird of paradise allows
him to put on a spectacular courtship dance to attract fe-
males and to give her a basis on which to select a suitable
mate (see Fig. 32.5). Defense and courtship displays are ex-
aggerated and are always performed in the same way so
that their meaning is clear.
Tactile Communication
Tactile communication occurs when one animal touches an-
other. For example, gull chicks peck at the parents beak in
order to induce the parent to feed them (see Fig. 32.3). A
male leopard nuzzles the females neck to calm her and to
stimulate her willingness to mate. In primates, grooming
one animal cleaning the coat and skin of anotherhelps
cement social bonds within a group.
Honeybees use a combination of communication meth-
ods, but especially tactile, to impart information about the
environment. When a foraging bee returns to the hive, it per-
forms a waggle dance that indicates the distance and the di-
rection of a food source (Fig. 32.11). As the bee moves
between the two loops of a gure 8, it buzzes noisily and
shakes its entire body in so-called waggles. Outside the hive,
the dance is done on a horizontal surface, and the straight
run indicates the direction of the food. Inside the hive, the
angle of the straight run to that of the direction of gravity is
the same as the angle of the food source to the sun. In other
words, a 40 angle to the left of vertical means that food is
40 to the left of the sun. Bees can use the sun as a compass
to located food because their biological clocks, as discussed
in the reading on page 682, allow them to compensate for the
movement of the sun in the sky.
Animals use a number of different ways to
communicate, and communication facilitates
cooperation.
Figure 32.11 Communication among bees.
a. Honeybees do a waggle dance to indicate the direction of food. b. If the dance is done outside the hive on a horizontal surface, the straight run
of the dance will point to the food source. If the dance is done inside the hive on a vertical surface, the angle of the straightaway to that of the
direction of gravity is the same as the angle of the food source to the sun.
Direction of flower
a. Waggle dance b. Components of dance
Certain behaviors in animals recur at regular intervals. Behav-
iors that occur on a daily basis are said to have a circadian
(about a day) rhythm. For example, some animals, like humans,
are usually active during the day and sleep at night. Others,
such as bats, sleep during the day and hunt at night. There are
also behaviors that occur on a yearly basis. For example, in the
Northern Hemisphere, birds migrate south in the fall, and the
young of many animals are born in the spring. Such behaviors
have a circannual rhythm.
There were two hypotheses regarding the control of circa-
dian rhythms: either their timing is controlled externally, or
there is an internal timing device, often called a biological clock
(Fig. 32B). These alternative hypotheses have been tested in
crickets, which regularly call every night to attract females.
When laboratory crickets are kept in a room under constant
conditions with lights continually on or continually off, they
continue to call every night, only calling starts as much as 26
hours later than it did on the day before. But exposure to night
and day cycles will right the cycles; therefore, it was possible to
conclude that there is a biological clock, but it does not keep per-
fect time and must be reset by environmental stimuli. Similar re-
sults have been obtained with all sorts of animals, from ddler
crabs to humans.
At a minimum, a circadian system must have three compo-
nents (Fig. 32B). There must be a means to reset a pacemaker ac-
cording to the current environmental light-dark cycle; a
biological clock that keeps time; and the rhythmic behavior it-
self. By now, there is a general consensus that the mammalian
biological clock is a collection of nerve cells in the hypothalamus
of the brain known as the suprachiasmatic nucleus (SCN). Elec-
trical and drug stimulation of the SCN upsets circadian rhythms
and destruction of the nucleus does away with many rhythmic
behaviors. Investigators have also discovered a number of
genes whose protein products control the activity of the clock.
Less is known about the other components of the circadian sys-
tem. Whether the receptor that resets the clock is in the eye or
not is still to be determined. In one recent study, investigators
found that exposing the back of a knee to light reset the clock
judged by body temperature and hormone levels. They con-
cluded that any portion of the skin contains photoreceptor pro-
teins that communicate with and reset the biological clock.
A number of investigators are interested in the observation
that environmental light suppresses melatonin production by
the pineal gland while we are awake. Conversely, as it gets dark,
melatonin levels rise and we get sleepy. This knowledge causes
some people to take melatonin for the symptoms of insomnia
and jet lag. When we travel by airplane from one part of the
world to another it is difcult for our circadian system to adjust,
and symptoms like insomnia, fatigue, headache, gastrointesti-
nal distress, and moodiness, occur. A medication for these
symptoms would also be helpful to one out of every ve people
whose work shifts between day and night. Jet lag symptoms are
reduced by exposure to daylight in the afternoon after west-
ward ights and in the early morning after eastward ights.
Similarly, an adjustment to a nighttime work shift is enhanced
by exposure to bright light at night and to darkness during the
day. Because light wakes us up and melatonin makes us sleepy,
their use at different times of the day may be helpful in resetting
the biological clock.
Many people are intrigued by the idea that some psychiatric
conditions might be due to a disorder of the circadian system.
The best example is seasonal affective disorder (SAD) which af-
fects as much as 5% of the general population and is character-
ized by depressions during the fall and winter. The hypothesis
that short winter days are responsible for SAD has led to suc-
cessful therapies based on exposure to bright light. Circadian
rhythms are clinically signicant in other ways. The incidence of
heart attacks, sudden cardiac death, and stroke peak in the late
morning, and certain cancer drugs are more effective when
given in the day or night. This is an exciting time for research
concerning the human circadian system especially because such
research is expected to have important applications that will
help control many human ills, from those that are quite serious
to those that are simply a nuisance.
682
Biological Rhythms
0
Time
H
o
r
m
o
n
e

L
e
v
e
l
Internal
biological
clock
keeps time.
resetting
External
light/dark
periods
reset
clock.
Hormone
level varies
with time
of day.
Figure 32B A circadian system.
A circadian system has three components, as shown.
Altruism Versus Self-Interest
Altruism is behavior that has the potential to decrease the
lifetime reproductive success of the altruist while beneting
the reproductive success of another member of the group.
In insect societies especially, reproduction is limited to only
one pair, the queen and her mate. For example, among
army ants the queen is inseminated only during her nuptial
ight, and thereafter she spends her time reproducing. The
society has three different sizes of sterile female workers.
The smallest workers (3 mm), called the nurses, take care of
the queen and larvae, feeding them and keeping them
clean. The intermediately-sized workers, constituting most
of the population, go out on raids to collect food. The sol-
diers (14 mm), with huge heads and powerful jaws, run
along the sides and rear of raiding parties where they can
best attack any intruders.
Can the altruistic behavior of sterile workers be ex-
plained in terms of reproductive success? A given gene can
be passed from one generation to the next in two quite dif-
ferent ways. The rst way is direct: a parent can pass the
gene directly to an offspring. The second way is indirect: an
animal can help a relative reproduce and thereby pass the
gene to the next generation via this relative. Direct selection
is natural selection that can result in adaptation to the envi-
ronment when the reproductive success of individuals dif-
fers. Indirect selection is natural selection that can result in
adaptation to the environment when individuals differ in
their effects on the reproductive success of relatives. The in-
clusive tness of an individual includes personal reproduc-
tion and reproduction of relatives.
Among social bees, social wasps, and ants, the queen is
diploid but her mate is haploid. If the queen has had only
one mate, sister workers are more closely related to each
other (sharing on average 75% of their genes) than they are
to their potential offspring (with which they would share on
average only 50% of their genes). Therefore, a worker can
achieve a greater inclusive tness benet by aiding her
mother (the queen) to produce additional sisters than by di-
rectly reproducing herself. Under these circumstances, be-
havior that appears to be altruistic is more likely to evolve.
Indirect selection can also occur among animals whose
offspring receive only a half set of genes from both parents.
Consider that your brother or sister shares 50% of your
genes, your niece or nephew shares 25%, and so forth. This
means that the survival of two nieces (or nephews) is worth
the survival of one sibling, assuming they both go on to re-
produce.
Among chimpanzees in Africa, a female in estrus fre-
quently copulates with several members of the same group,
and the males make no attempt to interfere with each others
matings. How can they be acting in their own self-interest?
Genetic relatedness appears to underlie their apparent altru-
ism; members of a group share more than 50% of their genes
in common because members never leave the territory in
which they are born.
32.5 Sociobiology and Animal
Behavior
Sociobiology applies the principles of evolutionary biology
to the study of social behavior in animals. Sociobiologists de-
velop hypotheses about social living based on the assump-
tion that a social individual derives more reproductive
benets than costs from living in a society. Then they perform
a cost-benet analysis to see if their hypotheses are correct.
Group living does have benets under certain circum-
stances. It can help an animal avoid predators, rear off-
spring, and nd food. A group of impalas is more likely to
hear an approaching predator than a solitary one. Many sh
moving rapidly in many directions might distract a would-
be predator.
Pair bonding of trumpet manucodes helps the birds
raise their young. Due to their particular food source, the fe-
male cannot rear as many offspring alone as she can with the
males help. Weaver birds form giant colonies that help pro-
tect them from predators, but the birds may also share infor-
mation about food sources. Primate members of the same
troop signal to one another when they have found an espe-
cially bountiful fruit tree. Lions working together are able to
capture large prey, such as zebra and buffalo.
Group living also has its disadvantages. When animals
are crowded together into a small area, disputes can arise
over access to the best feeding places and sleeping sites.
Dominance hierarchies are one way to apportion resources,
but this puts subordinates at a disadvantage. Among red
deer, the ability of a hind to rear sons is dependent on her
dominance. Only large, dominant females can successfully
rear sons; small subordinate females tend to rear daughters.
From an evolutionary point of view, sons are preferable be-
cause, as a harem master, sons will result in a greater num-
ber of grandchildren. However, sons, which tend to be
larger than daughters, need to be nursed more frequently
and for a longer period of time. Subordinate females do not
have access to enough food resources to adequately nurse
sons and, therefore, they tend to rear daughters and not
sons. Still, like the subordinate males in a baboon troop, sub-
ordinate females in a red deer harem may be better off in t-
ness terms if they stay with a group, despite the cost
involved.
Living in close quarters means that illness and parasites
can pass from one animal to another more rapidly. Baboons
and other types of social primates invest much time in
grooming one another, and this most likely helps them re-
main healthy.
Social living has both advantages and
disadvantages. Only if the benets, in terms of
individual reproductive success, outweigh the
disadvantages will societies evolve.
Helpers at the Nest
In some bird species, offspring from one clutch of eggs may
stay at the nest helping parents rear the next batch of off-
spring. In a study of Florida scrub jays, the number of edg-
lings produced by an adult pair doubled when they had
helpers. Mammalian offspring are also observed to help
their parents (Fig. 32.12). Among jackals in Africa, pairs
alone managed to rear an average of 1.4 pups, whereas pairs
with helpers reared 3.6 pups.
Is the reproductive success of the helpers increased by
their altruistic behavior? It could be if the chance of their re-
producing on their own is limited. A ock of green wood-
hoopoes (Phoeniculus purpurens), an insect-eating bird of
Africa, may have as many as sixteen members but only one
breeding pair. The other sexually mature members help feed
and protect the edglings and protect the home territory
from invasion by other green wood-hoopoes. Resources are
limited, particularly because nest sites are rare. Few acacia
trees have suitable cavities to serve as nest sites and those
that do are often occupied by other species. Moreover, pre-
dation by snakes within the cavities can be intense; even if a
pair of birds acquire an appropriate cavity, they would be
unable to protect their offspring by themselves. Therefore,
the cost of trying to establish a territory is clearly very high.
What are the benets of staying behind to help? First, a
helper is contributing to the survival of its own kin. Therefore,
the helper actually gains a tness benet (albeit a smaller ben-
et than it would achieve were it a breeder). Second, a helper
is more likely than a nonhelper to inherit a parental
territoryincluding other helpers. Helping, then, involves
making a minimal, short-term reproductive sacrice in order
to maximize future reproductive potential. Once again, an ap-
parently altruistic behavior turns out to be an adaptation.
Inclusive tness is measured by genes an
individual contributes to the next generation, either
directly by offspring or indirectly by way of
relatives. Many of the behaviors once thought to
be altruistic turn out, on closer examination, to be
examples of kin selection, and are adaptive.
I
s it ethical to keep animals in zoos where
they are not free to behave as they do in
the wild? If we keep animals in zoos are
we depriving them of their freedom?
Some point out that freedom is never ab-
solute. Even an animal in the wild is re-
stricted in various ways by its abiotic and
biotic environment. The so-called ve free-
doms are to be free of starvation, cold, in-
jury, and fear, as well as free to wander
and express ones natural behavior. Per-
haps its worth giving up a bit of the last
freedom to achieve the rst four? Many
modern zoos do keep animals in habitats
that nearly match their natural one so that
they do have some freedom to roam and
behave naturally. Perhaps, too, we should
consider the education and enjoyment of
the many thousands of human visitors to a
zoo compared to the loss to a much
smaller number of animals kept in a zoo?
Today, reputable zoos rarely go out
and capture animals in the wildthey
usually get their animals from other zoos.
Most people feel it is never a good idea to
take animals from the wild except for very
serious reasons. Certainly, zoos should not
be involved in the commercial and often il-
legal trade in wild animals which still goes
on today. When animals are captured it
should be done by skilled biologists or
naturalists who know how to care and
transport the animal.
Many zoos today are involved in the
conservation of animals. They provide the
best home possible while animals are re-
covering from injury or their numbers are
increased until they can be released to the
wild. Perhaps we can look upon zoos fa-
vorably if they can show that their animals
are being kept under good conditions, and
that they are also involved in the preserva-
tion of animals.
Questions
1. Do you think it is ethical to keep animals
in zoos? Under particular circumstances?
Explain.
2. Do the animals that are descended from
zoo animals have the right to be protected
as their parents were? Why or why not?
3. Do the same concerns about zoos also
apply to aquariums? Why or why not?
Figure 32.12 Inclusive tness.
A meerkat is acting as a baby-sitter for its young sisters and brothers
while their mother is away. Could this helpful behavior contribute to
the baby-sitters inclusive tness?
Go To Student OLC
Summarizing the Concepts
Ethologists study how animals are organized to perform a particular
behavior and how it benets them to survive and reproduce.
Hybrid studies with lovebirds produce results consistent with the hy-
pothesis that behavior has a genetic basis. Garter snake experiments in-
dicate that the nervous system controls behavior. Aplysia DNAstudies
indicate that the endocrine system also controls behavior.
The environment is inuential in the development of behavioral re-
sponses, as exemplied by an improvement in laughing gull chick
begging behavior and an increased ability of chicks to recognize par-
ents. Modern studies suggest that most behaviors improve with expe-
rience. Even behaviors that were formerly thought to be xed action
patterns (FAPs) or otherwise thought to be inexible sometimes can be
modied.
Song learning in birds involves various elementsincluding the
existence of a sensitive period when an animal is primed to learnand
the effect of social interactions.
Traits that promote reproductive success are expected to be advan-
tageous overall despite any possible disadvantage. Males who pro-
duce many sperm are expected to compete to inseminate females.
Females who produce few eggs are expected to be selective about
their mates. Experiments with satin bowerbirds and birds of par-
adise support these bases for sexual selection.
The raggiana bird of paradise males gather in a lekmost likely
because females are widely scattered, as is their customary food source.
The food source for a related species, the trumpet manucode, is readily
available but less nutritious. These birds are monogamousit takes
two parents to rear the young.
A cost-benet analysis can be applied to competition between
males for mates in reference to a dominance hierarchy (e.g., baboons)
and territoriality (e.g., red deer).
Animals that form social groups communicate with one another. Com-
munications such as chemical, auditory, visual, and tactile signals fos-
ter cooperation that benets both the sender and the receiver.
There are benets and costs to living in a social group. If animals
live in a social group, it is expected that the advantages (e.g., help to
avoid predators, raise young, and nd food) will outweigh the disad-
vantages (e.g., tension between members, spread of illness and para-
sites, and reduced reproductive potential). This expectation can
sometimes be tested.
32.5 Sociobiology and Animal Behavior
In most instances, the individuals of a society act to increase their own
reproductive success. Sometimes animals perform altruistic acts, as
when individuals help their parents rear siblings. There is a benet to
this behavior when one considers inclusive tness, which involves
both direct selection and indirect selection.
In social insects, altruism is extreme but can be explained on the
basis that the insects are helping a reproducing sibling survive. Astudy
of wood-hoopoes, an African bird, shows that younger siblings may
help older siblings who reared them until the younger get a chance to
reproduce themselves.
Studying the Concepts
1. What two aspects of behavior particularly interest etholo-
gists? 672
2. Describe an experiment with lovebirds, and explain how it
shows that behavior has a genetic basis. 672
3. An experiment with garter snakes showed that what system
is involved in behavior? Explain the experiment and the
results. 67273
4. Studies of Aplysia DNAshow that the endocrine system is
also involved in behavior. Explain. 673
5. Some behaviors require practice before developing
completely. How does the experiment with laughing gull
chicks support this statement? 674
6. An argument can be made that social contact is an important
element in learning. Explain this with reference to imprinting
in mallard ducks and song learning in white-crowned spar-
rows. 675
7. Why would you expect behavior to be subject to natural
selection and be adaptive? 676
8. Reproductive behavior sometimes seems tied to how an
animal acquires food. Explain with reference to the two bird
of paradise species discussed in this chapter. 676
9. Explain how the anatomy and behavior of dominant male
baboons is both a benet and a drawback. 677
10. Give examples of the different types of communication
among members of a social group. 68081
11. What is a cost-benet analysis and how does it apply to liv-
ing in a social group? Give examples. 683
12. How can altruism, as dened on page 683, be explained on
the basis of self-interest? 68384
Testing Yourself
Choose the best answer for each question.
1. Which question is least likely to interest a behaviorist?
a. How do genes control the development of the nervous
system?
b. Why do animals living in the tundra have white coats?
c. Does aggression have a genetic basis?
d. Why do some animals feed in groups and others feed singly?
e. Behaviorists only study specic animals.
2. Female sage grouse are widely scattered throughout the
prairie. Which of these would you expect?
a. Amale will maintain a territory large enough to contain at
least one female.
b. Male and female birds will be monogamous, and both will
help feed the young.
c. Males will form a lek where females will choose a mate.
d. Males will form a dominance hierarchy for the purpose of
distributing resources.
e. All of these are correct.
3. White-crowned sparrows from two different areas sing with a
different dialect. If the behavior is primarily genetic, newly
hatched birds from each area will
a. sing with their own dialect.
b. need tutors in order to sing in their dialect.
c. sing only when a female is nearby.
d. learn to sing later.
e. Both a and c are correct.
4. Orangutans are solitary but territorial. This would mean
orangutans defend their territorys boundaries against
a. other male orangutans.
b. female orangutans.
c. all types of animals, whether orangutans or not.
d. animals that prey on them.
e. Both a and b are correct.
5. The resplendent plumes of a raggiana bird of paradise are
due to the fact that birds with the best display
a. are dominant over other birds.
b. have the best territories.
c. are chosen by females as mates.
d. are chosen by males and females as companions.
6. Subordinate females in a baboon troop do not produce off-
spring as often as dominant females. It is clear that
a. the cost of being in the troop is too high.
b. the dominant males do not mate with subordinate females.
c. subordinate females must benet in some way from being
in the troop.
d. Subordinate females should leave the troop.
e. Both a and b are correct.
7. German blackcaps migrate southeast to Africa, and Austrian
blackcaps y southwest to Africa. The fact that hybrids of
these two are intermediate shows that
a. the trait is controlled by the nervous system.
b. nesting is controlled by hormones.
c. the behavior is at least partially genetic.
d. behavior is according to the sex of animals.
8. At rst laughing gull chicks peck at any model that looks like
a red beak; later they will not peck at any model that does not
look like a parent. This shows that the behavior
a. is a xed action pattern.
b. undergoes development after birth.
c. is controlled by the nervous system.
d. is under hormonal control.
9. Which answer is based on anatomy? Males compete because
a. they have the size and weapons with which to compete.
b. they produce many sperm for a long time.
c. the testes produce the hormone testosterone.
d. only then do females respond to them.
10. Which answer is in keeping with evolutionary theory?
Females are choosy because
a. they do not have the size and weapons with which to
compete.
b. they invest heavily in the offspring they produce.
c. ovaries produce the hormones estrogen and progesterone.
d. they need time to get ready to respond.
Thinking Scientically
1. In an experiment to determine the control of behavior, young
guinea pigs are placed in a cage and are fed a type of food
they do not ordinarily eat in the wild. When released, the
guinea pigs are offered various types of food, including the
one type they were fed while caged (page 673).
a. How might you attempt to show that the nervous system
is involved in food gathering and eating?
b. What will you conclude if all groups of guinea pigs choose
only the type of food they were recently fed?
c. What will you conclude if the guinea pigs choose only the
type of food they ordinarily eat in the wild?
2. Rewarding an animal with food helps them learn a behavior.
Based on this nding (page 674),
a. what advice might you have for parents?
b. what part of the brain must be involved in learning the
behavior?
c. what other experiments might support your hypothesis?
Understanding the Terms
altruism 683
behavior 672
communication 680
dominance hierarchy 677
imprinting 675
inclusive tness 683
learning 674
operant conditioning 674
pheromone 680
sexual selection 676
society 680
sociobiology 683
territoriality 679
Match the terms to these denitions:
a. Behavior related to the act of marking or defend-
ing a particular area against invasion by another species
member; area often used for the purpose of feeding, mating,
and caring for young.
b. Social interaction that has the potential to
decrease the lifetime reproductive success of the member
exhibiting the behavior.
c. Signal by a sender that inuences the behavior of
a receiver.
d. Chemical substance secreted by one organism that
inuences the behavior of another.
e. Increase in reproduction that results from direct
selection and indirect selection.
Using Technology
Your study of animal behavior is supported by these
available technologies.
Essential Study Partner CD-ROM
Ecology Behavior
Visit the Mader web site for related ESP activities.
Exploring the Internet
The Mader Home Page provides resources and
tools as you study this chapter.
http://www.mhhe.com/biosci/genbio/mader

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669

Ecology began as a descriptive

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