THE ETHOGRAM ANIMAL BEHAVIOR RESEARCH and Supported by a Science Education Partnership Award from the National Center for Research Resources, National Institutes of Health Janet Crews Stan Braude Carol Stephenson Terrilyn Clardy science curriculum for grades 5-8
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T H E
ETHOGRAMANIMAL BEHAVIOR RESEARCH
and
Supported by a Science Education
Partnership Award from the National
Center for Research Resources,
National Institutes of Health
Janet CrewsStan Braude Carol StephensonTerrilyn Clardy
science curriculum for grades 5-8
science curriculum for grades 5-8
Supported by a Science Education
Partnership Award from the National
Center for Research Resources,
National Institutes of Health
Janet CrewsStan Braude Carol StephensonTerrilyn Clardy
T H E
ETHOGRAMANIMAL BEHAVIOR RESEARCH
and
2
Authors: Janet Crews, Wydown Middle School, Clayton School District
Stan Braude, Department of Biology, Washington University
Carol Stephenson, Saint Louis Zoo
Terrilyn Clardy, Sumner High School, St. Louis Public Schools
Animal behavior films produced by Rosie Koch, BAO Productions
This curriculum was developed by A Partnership Linking Formal and Informal Education, fundedby Science Education Partnership Award R25 RR 15603 from the National Center for ResearchResources, National Institutes of Health. Principal Investigator: Ralph S. Quatrano, Professor of Biology, Washington UniversityProject Director: Victoria L. May, Director of Science Outreach, Washington University Evaluator: Jennifer L. Heim, St. Louis Science Center
Washington University Science OutreachOne Brookings Drive, Campus Box 1137, St. Louis, MO 63130(314) 935-6846, www.so.wustl.edu
3
C O N T E N T S
S E C T I O N 1 Introduction and Overview of the Curriculum . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
S E C T I O N 2 The Ethogram: an Introduction to the Study of Animal Behavior . . . . . . . . . . . . 13
S E C T I O N 3 Introduction to Scientific Inquiry and Sampling Methods and Data Analysis . . . 25
S E C T I O N 4 Animal Behavior Research Projects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
S E C T I O N 5 Appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
INTRODUCTION AND OVERVIEW OF THEANIMAL BEHAVIOR CURRICULUM
5
7
The study of animal behavior is important for basic
understanding of the world around us, and has practical
applications for conservation, agriculture, and medicine. The
study of animal behavior in the classroom also provides
opportunities for teaching students about the process of
doing science, as well as developing basic skills of
observation, description, and quantification. It helps teachers
address components of all four of the Missouri Show Me
performance standards and includes content standards from
science, social studies and communication arts.
In this unit, students, will create one of the basic descriptive
tools of animal behavior, the ethogram. A complete
ethogram is a dictionary of all of the behaviors that a
particular species performs. Biologists publish ethograms of
their study species so that they can communicate with each
other about the behaviors they investigate. Generating their
own ethograms will give students multiple opportunities to
hone their skills of observation, description and
quantification. Students will make preliminary ethograms in
the classroom and at home. They will then generate an
ethogram of one exotic species at the zoo by viewing a
video of that animal. They will then use their ethograms in
completing a research project at the zoo.
This unit uses the attraction of studying exotic species at the
zoo to teach students about the process of doing scientific
research. They will explore one zoo species in depth and
then write up a research proposal.
While this unit is designed to utilize the resources of the zoo,it can be completed by using a local park, class study area oncampus, or even using a class pet. Financial resources shouldnot keep a teacher from using this curriculum.
Teachers have the option of planning one, two, or three zoo
visits. See graph Zoo visit options for Section 4, page 11.
Two visits are ideal for this unit. The preliminary zoo trip,
if taken, gives students an opportunity to make
observations that can help guide development of their
scientific research proposals.
In the second, third (or only) visit to the zoo, students will
take data that will help them test a hypothesis that follows
from a scientific question about their chosen study species
at the zoo. Depending on time available, the class may
spend part of the day visiting the sites of each study.
Students can tell their classmates about their chosen
study animal.
It is the intent of this curriculum that teachers will use
the zoo and other local resources, not just for entertainment
field trips, but for powerful field classes. The power of a
focused learning experience, coupled with intense pre- and
post-field work in the classroom, is what increases student
achievement. It is important for the teacher to wisely
decide how this curriculum can best support existing
district curriculum. Does it help you teach scientific
method? Taxonomy? Explore native species? Or is it best
suited for cross-curricular projects? Can it help you provide
a rationale for a service learning project? The choice is up
to the teacher.
Finally, it is also up to the teacher to select the best time of
the year to implement this curriculum. Testers suggest the
spring, as you have had time to work with your students,
and they have matured in their skills. It is also more
appropriate weather for outdoor field classes.
Whatever your situation, this curriculum is yours to adapt
and incorporate to help you better teach and engage your
students. The use of ethograms truly lets the students (and
teacher) feel like real scientists.
1.1 INTRODUCTION
8
S E C T I O N 2 THE ETHOGRAM: AN INTRODUCTION TO THE STUDY OF ANIMAL BEHAVIOR. . . . 13
— Empty aquarium with bedding and rodent supplies.
Tape the bottom edge of the cage and label each
corner with letters A–D, creating quadrants.
— Stopwatch
— All Occurrences Sampling data sheet
— Scan Sampling data sheet: Habitat use
— Scan Sampling data sheet: Marked individuals
— Focal Sampling data sheet
— Looking for Patterns handout (optional)
— Which Sampling Method Works Best? worksheet
3.1 WHICH SAMPLING METHOD WORKS BEST?
27
Procedure
1 Ask students how we could find out how middleschool students spend their time. Write possibleanswers on the board.
2 Ask students what a time budget might be. Writepossible answers on the board.
3 Ask them what a time budget for a middle school studentmight look like. As a class, design one on the board.
4 Hand out the All Occurrences Sampling data sheet.Discuss how to tally. What does all occurrences mean?What does tally mean? What is a focal behavior? Referto key terms. Why are the behaviors defined? Discussany unclear parts.
5 Position the students in a circle with the rodent cage inthe center on a desk. Start the time and have studentstally all occurrences of behavior for 5-7 minutes.
6 Have students total the tallies. To find the percentage,divide the total of each behavior by the total of alloccurrences, then multiply by 100 (e.g., 5 sniffs of fooddivided by 52 total behavior occurrences multiplied by 100 = 9.6% of the time spent sniffing food). Discuss problems or questions. Many students find this part difficult.
7 Hand out the two Scan Sampling data sheets. Discusshow to tally on these sheet. What does scan mean?What is a quadrant? What are intervals? What area isA? B? C? D? Refer to key terms.
8 Again, group the students in a circle. Place the quadrantlabeled cage into the center of the circle. Move therodent into the new cage. Begin the time as soon as it isinside. Speak or ring a bell every 20 seconds to remindstudents to tally data. Students can work in cooperativegroups with a timekeeper, an observer who calls out theobservations, a recorder who writes, and one who takescare of the materials and sets up the quadrants.
9 Ask students to total tallies and find percentages.Discuss problems.
10 Have students look over their data and see if they caninfer anything about rodents from the data theycollected. What is missing? Can any clearunderstanding or answer come from these twosamplings of data? Does it need to be compared tosomething else?
11 Point out the different questions on the worksheets.How is each type of sampling suited to each question?
12 Explain that all animals are engaging in some type ofbehavior all the time –- even if it doesn’t look like itand only one kind of behavior at a time.
13 Scientists use two terms to separate types ofbehaviors — states and events. Write these terms anddefinitions on the board or overhead.
14 Have students look at the two different Scan Samplingdata sheets. Are the behaviors listed mostly states orevents? (These are mostly events. Eating andgrooming can be states — depending on length oftime) What if the rodent was sleeping? Would this type of sampling method be helpful? (not for only 5 minutes — but it would for longer periods of time)
15 Now look at the All Occurrences Sampling data sheet.Are you recording states or events? (This should bemostly states – it is likely that an event could be lost ifit is between recording times. Ask students if this wasobserved by anyone.)
16 How do scientists decide which type of ethogram to use? (It is determined by the question they seek to answer.)
17 Hand out Which Sampling Method Works Best? or write the questions on the board. As a group, discusswhich sampling methods would provide the mostuseful information in answering the questions, orassign the worksheet as homework.
18 Have students summarize their understanding ofeach sampling type in their science journals.
WHICH SAMPLING METHOD WORKS BEST?3.1
28
Teacher Tips
SAMPLING METHOD DEFINITION CONSIDERATIONS
All Occurrences Recording each time a Especially useful for studying rare or infrequent
Sampling behavior is performed, either behaviors. Also useful if you are interested in
on a focal animal or among whether a behavior occurs more in one setting, by
all animals present. one sex, by one age class, etc. Time budgets cannot
be derived from this type of data. Not useful if
you are interested in the amount of time spent in a
given state; more useful in studying behavioral events.
Focal Animal Collecting data focusing Video is a useful tool for this type of sampling.
Sampling on one individual animal Working in teams is also useful. This is the only type
at a time, noting every of sampling that gives us data with which we can
behavior that animal generate a flow chart. Time budgets can be derived
performs, and then pooling from this type of data. Especially useful if you are
the data for a number of interested in the amount of time spent in a given state;
different animals. less appropriate if you are studying behavioral events.
Scan Sampling Collecting data at specified Still photos may be useful in doing this sampling.
time intervals, and noting This method is useful in asking question about habitat
what each animal is preference, food preference, or preferred nearest
currently doing or where neighbor. Time budgets can be derived from this type
they are located. of data. The fraction of time spent in a given state
can be derived from this data. If you are studying
behavioral events, you are likely to miss them with
this sampling method.
Different sampling methods can be used to answer the same question. It can be an interesting
exercise for advanced students to compare the answer they get from collecting observations with
different sampling methods.
WHICH SAMPLING METHOD WORKS BEST?3.1
All Occurrences: Recording each time a behavior is performed, either on a focal animal or
among all animals present.
Uses: Especially useful for studying rare of infrequent behaviors. Also useful if you are
interested in whether a behavior occurs more in one setting, by one sex, by one age class,
etc. Time budgets cannot be derived from this type of data. Not useful if you are interested
in the amount of time spent in a given state; more useful in studying behavioral events.
What activity does a classroom animal spend most of its time doing?
Provides insufficient informationfor two or all of the following –
conclusion, inferences,recommendations
1 point does not meet
Student did not perform theexperiment honestly and/or
accurately1 point does not meet
Does not meet = Below 18 points
A L T E R N A T I V E S C O R I N G G U I D E : G R O U P M E T H O D
ANALYSIS OF RESULTS AND WRITE UP OF RESEARCH PROJECT4.10
78
Lesson Overview
Students have the opportunity to reflect over their
conclusions and what they have learned during this
project. This could also include students evaluating their
peers’ presentations.
Student Objectives
After completing this activity, students will be able to:
1 answer reflective questions on various aspects of
research they conducted.
2 compare and contrast their prior knowledge of animal
behavior research with their post-research knowledge.
Timeline
1 class period
Procedure
1 Have students share their projects with each other. This
could be for the whole class or in like-species groups.
2 Ask students to record questions, comments and
compliments about their peers’ projects.
3 Allow time for students to comment after each
presentation.
4 Hand out Student Self Reflection.
5 Ask the students to demonstrate what they have
learned by answering the questions as completely as
they can. Encourage them to use specific examples
from their research.
6 Have the students compare what they knew when
first started doing animal observations and what they
know now. Have their knowledge and understanding
increased?
4.11 REFLECTION / EVALUATION
79
1 What causes behavior?
2 How do scientists measure, test, or explain behavior?
3 How certain do you feel that the conclusion you reached regarding behavior of
your animal is accurate? Why?
4 Is your research project repeatable by any other scientists, just from using your
report? Why or why not?
5 What did you see in other projects that you would add to your own? Why?
nameACTIVITY
STUDENT SELF-REFLECTION
4.11
80
Lesson Overview
Students or teachers may choose to continue with this
topic in a variety of extension activities.
Student Objectives
Students will continue to expand their knowledge
of behavioral studies by completing a deeper
investigation.
Timeline
Varies. Based on the project chosen.
Procedure
1 Either give the students choices to select from, ask
them to brainstorm topics of interest, or select one of
your choice to proceed with.
2 Suggested extensions include:
• Just as the zoo provided you with a video on your
species, produce your own video/hyperstudio on your
species, or one of your choosing. Will yours be used to
practice ethogram techniques? Or will it be like a
National Geographic Explorer Report? If no technology
is available, create a written natural history of zoo
species like the ones in this unit. If each student
creates one, you might have the whole zoo covered!
• Did your project raise more questions about your
animal’s behavior, or about a similar animal? If so, use
your scientific question to develop a science fair project.
Submit it to your school or community’s science fair!
• How well can your students apply their ethogram skills
to other species? Show the monkey and/or baboon tape
from the first lesson again. This time, ask the students
to write a question, make a hypothesis and create an
appropriate ethogram for the animal. Then show it
again and let them collect their data. Have them write a
conclusion. This could serve as a final exam.
• Use your new found knowledge to do a behavioral study
of an animal close to home. Study a classroom pet
(hamsters, fish, lizards), or one from your school yard
(birds, squirrels, rolypolys, worms, ants) or one from a
nearby park. You could even study other students in
your school. Try to solve a problem that currently exists
that is behavioral in nature. Use the same process.
• Justify a behavior change in your school or community
(service project) by collecting human behavior data.
Examples: recycling issues, SUV usage, test out a
proverb or saying you’ve heard.
• Share what you have learned with other experts. Invite
zoo personnel to your school, or go there, and share
your findings. If you do a study of your school, school
ground or local park, invite concerned parties to your
presentations. Use this scientific methodology to solve a
problem that affects all citizens.
Make your own extension! Be creative!
4.12 FURTHER INVESTIGATIONS
S E C T I O N 5
APPENDICES
81
82
CRICKETS IN THE CLASSROOM
Field crickets are excellent subjects for studying behavior in
the classroom because they have a wide repertoire of social
behaviors. However, in order to see these behaviors we need
to observe the right crickets in the right environment.
Age
Many pet stores sell large and small crickets, but size
does not guarantee maturity. Juveniles will not engage
in territorial, courtship, or oviposition behaviors.
Juveniles, even large juveniles, do not have wings; they
merely have wing buds or pads. Adults have dark hard
forewings that the males use to produce their songs.
Comparisons between juveniles and adults would be
an interesting project if you have enough crickets of
both age categories.
Sex
Males and females behave very differently. Males are
territorial and will sing to attract females. Males are
distinct in having thick veins in their forewings, which
produce their song when wings are moved past each
other. Females are most easily distinguished by their
ovipositors. This long thin organ extends from the tip of
the abdomen and is lowered into the soil where eggs
are deposited after the female copulates.
Housing
Males will defend a burrow against other males. If you
house your crickets in an aquarium, give each male a
small burrow to defend. (A toilet paper tube cut in
half and then lengthwise, or a large tampon tube, will
work well.) Males will behave differently in class
depending on their experience before class. Males
who are housed together will behave very differently
than males who are isolated until class. Isolated males
will scramble to find and then defend the new burrows.
Females will lay eggs in moist (not soaking wet) sand or
loose soil. A petri dish can make a suitable oviposition
site. If a female is kept away from oviposition sites
before class, she is more likely to have eggs to lay
during class. She will probe the soil with the ovipositor
and deposit eggs if the humidity is appropriate. A
female isolated from males before class is also more
likely to respond to his courtship song during class.
Distractions
Crickets are more likely to sing at night (hence the
sound we associate with camping in the woods) but will
sing and behave during the daytime. Try to keep the
lights a bit dim and be very careful to avoid bumping the
table that the animal container is on. Crickets have their
acoustic organs (ears) on their forelegs and therefore
pick up vibrations from the surface they are standing on.
5.1 CARE T IPS FOR CLASSROOM ANIMALS
83
DO keep your mice clean. Your mouse cage should
be cleaned at least once a week. Be guided by your
nose. If you smell ammonia around the cage area, then
the time to change the bedding is past due. Put the
mice in a secure place, dump out the old bedding, wipe
out the bottom with a little soap and water, dry out
the cage, add fresh bedding, food, water and mice.
DO handle the mice yourself, DON’T delegate this
responsibility to members of the class. The mice will
respond by being much calmer when handled
regularly by the same person. Having you the teacher
handle and care for the mice is a requirement and
has the added benefit of reducing escapes and
nipped fingers.
DO keep your mice warm but not hot.
DON’T place in direct sunlight or in drafts from open
windows and air conditioners.
DO keep your mice dry. Sometimes water bottles leak
and flood the cage. Check your water bottle every day.
DO provide your mice with fresh drinking water. Water
bottles should be filled with fresh water twice weekly
and washed with a mild soap once a week.
DO’S AND DONT’S FOR MICE IN THE CLASSROOM
CARE T IPS FOR CLASSROOM ANIMALS5.1
84
DO make sure your mice have plenty of nutritious food.
The Rodent Lab Chow provided has everything
necessary to raise healthy litters of mice. Seeds and
other treats should be used sparingly for observations
of food preferences and foraging behaviors.
DON’T let the class decide what to feed the mice. Some
of their suggestions might be very good such as treats
of seeds, fresh vegetables and fruits, but others might
be harmful to the mice in the long run. Some foods
might be preferred by the mice but be nutritionally
incomplete.
DON’T trust the mice to eat what is best for them.
DO use the filter material provided for the top of the
cage. This material, when attached to the cage, acts as
a two way barrier by keeping unwanted objects out of
the cage and keeping dust and animal dander’s out of
your classroom.
DO use your mice as a resource and observational tool
for classroom discussion. Your class can learn a lot
about animal behavior by keeping a daily log of the
activities and behaviors shown by individual mice.
DO use the collected information from your class to
prepare an ethogram of mouse behavior.
DO limit the number of children observing the mice at
one time. Small groups of 3-4 work well.
DON’T allow children to tap on the cage to make the
mice active. Varying the time of day for observations
should expose the class to a range of behaviors.
DO use the mice for simple experiments that give the
mice a choice, such as; do they prefer cotton or kleenex
for their nest? do they prefer to build a nest in the open
or in a tube from a toilet paper roll?
DON’T use your mice as experimental subjects and
offer them objects that might be potentially harmful.
DO let your mice breed. Raising a litter of mouse pups
is a fascinating experience for the class. Mice have a
gestation period of 20-21 days. Try to clean the cage a
few days before the expected birth and then not again
for about a week after the female delivers. Most mice
choose a corner of the cage for urination. You should
clean these “wet spots” every 3-4 days to keep odors
under control. You may safely move the litter when they
are seven to ten days old for a more thorough cleaning.
DO remove the male and house him separately if you
don’t want another litter right away. The female will be
receptive to the male immediately after giving birth. If the
male remains, you’ll have another litter in about 3 weeks.
DO be careful when cleaning a cage containing young
mice of about 3 weeks. They are very excitable and
have amazing jumping abilities.
DON’T allow the cage to become overcrowded.
Overcrowding will be stressful to the mice and cause
aggression. A pecking order will be established by
fighting among the mice and the losers could
be severely injured. Overcrowding might also result in
cannibalism of new litters.
DON’T send the surplus mice home with members of
your class. Once out of the classroom you can’t control
what happens to the animals and their welfare may
be at stake.
DO bring surplus animals back to the university. We will
gladly take back progeny and parents when your
projects are finished. If you have mice from another
source, like a pet shop, ask if you can give them your
surplus mice.
CARE T IPS FOR CLASSROOM ANIMALS5.1
85
5.2 SAMPLE EXAMS
DIAGNOSTIC EXAM A
Ellie has been observing the mice in her classroom for
months. Last semester, Ellie noticed that the mice play with
each other like kittens or puppies. Ellie wondered why mice
play. Her friend David suggests that it is good exercise for the
mice and helps keep them fit. However Ellie has heard
that human play is an important part of learning how to
behave as an adult, and thinks that mice could play for
the same reason.
The adult female mouse in Ellie’s class is pregnant again and
Ellie wants to study the pups for her behavior research
project. Ellie thinks that the pups will behave differently than
the older mice since they are still learning how to behave.
Ellie has convinced her classmates to help her collect data
on the new mice but she has to organize her ideas and
organize the data collection so that it can be used to test
her hypothesis.
1 Clearly state Ellie’s research question.
2 Clearly state Ellie’s hypothesis.
3 What predictions has Ellie made that should be found if
her hypothesis is true?
4 Clearly state a prediction that Ellie and her classmates
can test by making observations of their class mice over
the next month.
5 Ellie’s classmates are going to help her collect data.
How can Ellie help make sure that the observations
recorded by different people are comparable and can
be pooled together?
6 Since the observers cannot watch every animal, all of
the time, explain how they can make their observations
so that they get the fairest test of their hypothesis?
7 Make a data sheet that Ellie’s classmates can use to
collect data for her.
8 Even before she has the data, we know what kind of
graph will best illustrate Ellie’s results. Draw the axes of
her graph and label the axes.
9 Fill in what you expect Ellie’s graph to look like if the
data supports her hypothesis (in red ink). Draw the
lines you expect to see if the data do not support Ellie’s
hypothesis (in blue ink).
86
KEY EXAM A
1 Why do mice play?
2 Mice play in order to better learn how to behave as adults.
3 Pups will behave differently than adults.
4 Pups will spend more of their time playing than adults. Or, pups will play less as they become adults.
5 Ellie can define what play looks like or teach her classmates what different behaviors look like so they
all call the same behavior the same thing.
6 They can watch each animal for the same amount of time each day.
7Animal’s Name Minutes observed playing Minutes observed not playing
Mother
Father
Pup 1
Pup 2Pup 3
8 and 9 Average time spent playing
adults pups age
or
SAMPLE EXAMS5.2
87
SAMPLE EXAMS5.2
DIAGNOSTIC EXAM B
David loves gray tree squirrels. He always has a bag of
peanuts in the pocket of his parka so that he can feed the
squirrels that he meets walking to or from school. He noticed
that some squirrels are very calm when they take his
peanuts and even sit next to him to eat the nuts. Others
seem very nervous and run away as soon as they get a nut.
David is curious why the squirrels behave so differently. His
friend Ellie suggests that the girl squirrels are larger and
braver and that the boy squirrels are the small, scared ones.
However, David is pretty sure that the same squirrels are
calm sometimes and nervous other times.
David has noticed that as soon as a dog comes in sight all
the squirrels run up the nearest tree. He thinks that the
squirrels might feel more secure when he is feeding them
near a tree. He has convinced his friends to help him collect
data to test his hypothesis. They agree to meet at Oak Knoll
Park on the following Sunday afternoon.
1 Clearly state David’s research question.
2 Clearly state David’s hypothesis.
3 What predictions can David make that should be found
IF his hypothesis is true?
4 Clearly state one prediction that David and his
classmates can test by making observations in Oak
Knoll Park on a Sunday afternoon.
5 David’s classmates are going to help him collect data.
How can David help make sure that the observations
recorded by different people are comparable and can
be pooled together?
6 Since the observers cannot watch every animal all of
the time explain how they can make their observations
so that they get the fairest test of their hypothesis.
7 Make a data sheet that David’s classmates can use to
collect data for him.
8 Even before he has the data, we know what kind of
graph will best illustrate David’s results. Draw the axes
of his graph and label the axes.
9 Fill in what you expect David’s graph to look like if the
data supports his hypothesis (in red ink). Draw the
lines you expect to see if the data do not support
David’s hypothesis (in blue ink).
88
SAMPLE EXAMS5.2
KEY EXAM B
1 Why are the squirrels sometimes calm and other times
nervous when they take his peanuts?
2 Squirrels feel safer when they are near a tree.
3 Squirrels will act more nervous and run away faster if
they are far from a tree.
4 The farther you are from a tree, the more quickly a
squirrel will take your peanut and run away.
5 David can define what he means by “run away”, “walk
around”, “beg for more” and other behaviors his friends
are likely to see. He can also make sure they offer the
same kind of peanuts and offer them in the same way.
6 They can test each squirrel at different distances from
the nearest tree. Or they can test each squirrel only once
but test different squirrels at different distances from the
nearest tree.
7
If one animal is tested at different distances, then
column 1 can be omitted and each friend gets a data
sheet for each animal.
8 and 9 Minutes spent taking and eating peanuts before
moving back to a tree.
Distance to Nearest Tree
Animal’s name Distance to Time squirrel Time squirrel moves Total minutes spent taking and eating or number nearest tree takes peanut back to tree peanuts before moving back to a tree.
89
SAMPLE OF A STUDENT REPORT
STUDENT SAMPLE
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STUDENT SAMPLE
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STUDENT SAMPLE
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STUDENT SAMPLE
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STUDENT SAMPLE
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STUDENT SAMPLE
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STUDENT SAMPLE
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?Scientists have long studied animal behavior in order to better understand them. From
anthropologists studying humans, to biologists observing animals in the wild, to
zoologists trying to help animals in captivity to survive, behavior is a source of knowledge
and evidence to scientists.
Zoos allow us to observe animals at a much closer level, just as class pets do in a classroom.
By gaining this understanding of animals in captivity, scientists can then look for those
behaviors in the wild, testing out theories and answering questions about the organisms.
Additionally, studying animal behavior helps scientists to predict how human actions will
affect animals and their environment in the future. Both are dependent on humans for
their continued existence. Humans have a responsibility to study animal behavior to
determine the needs of the animal kingdom and to determine when certain species are
being threatened. Human survival, as well as animal survival, depends on healthy
populations of diverse species.
Stan Braude, an animal behavior
researcher at Washington University,
has identified previously unknown
behaviors of the naked mole-rat.
Why Do Scientists Study Behavior?
he study of animal behavior is the
scientific study of everything that
animals do. Animals may be single-celled
organisms, invertebrates, fish, amphibians,
reptiles, birds, humans, or other mammals.
Animal behavior involves the investigation of
the relationship of animals to their physical
environment as well as to other organisms. It
includes topics such as how animals find
and defend resources, avoid predators,
choose mates and reproduce, and care for
their young.
People who study animal behavior are
typically trying to answer one or more of the
following four kinds of questions about
behavior. These questions are often referred
to as Tinbergen’s four questions after the
animal behaviorist Niko Tinbergen, who first
described them (see below).
T
Tinbergen’s Four Questions about Animal Behavior
WHYSanimal
1 What is the cause of the behavior?What mechanism is underlying it?What “triggered” it just now?
Answers may include both theexternal stimuli that affect behavior,and the internal hormonal andneural mechanisms that controlbehavior.
2 How did the behavior developwithin the individual’s lifetime?
Helps scientists learn how behaviorchanges over the lifetime of the ani-mal, and how these changes areaffected by both genes and experi-ence.
3 What function or functions does thebehavior serve?
Helps determine the behavior’seffect on the animal and its value inhelping the animal to survive orreproduce successfully in a particu-lar environment.
4 How did the behavior evolve over time?
Helps determine the origins ofbehavior patterns and how thesechange over generations.
from http://acunix.wheatonma.edu/kmorgan/AB_Careers/animal_behavior_careers.html
WHAT ISBEHAVIOR?
100
RESEARCHBEHAVIORAL
at the
ZOO STUDY
The zoo is developing a profile of typical behavior of mother and
infant pairs of antelopes during the first 30 days after birth. It will
be used as a baseline comparison of the species. The study
evaluates the behavioral interactions of the mother and infant that
are likely to be related to successful bonding and care. They include,
for the most part, measures of proximity, grooming and ursing.
In the wild, young hoofstock can be categorized either as
“hiders” or “followers,” depending on the species’ survival
strategy. Forest antelope species are known to hide their young
in the brush and return to nurse at very infrequent intervals
(sometimes as little as three times in a 24-hour period),
reducing the chance they are bringing the fawn to a predator’s
attention. In contrast, species from open grasslands give birth
to precocial young that are able to ”follow” their mothers.
Consequently, these young nurse much more frequently.
In the zoo’s commitment to leave infants with their mothers
whenever possible, it is in their best interest to understand what
is normal nursing behavior for each captive species. Knowing,
for example, that an individual species is a “hider,” animal care
staff will not be alarmed if they do not see a mother nursing her
young constantly.
Data for this study is collected via 24-hour video filming of the
exhibit, then tabulated by computer.
The Saint Louis Zoo applies animal
behavior research in its conservation
efforts both at the zoo and in the field.
The zoo’s efforts to breed threatened
and endangered animal species is
enhanced where knowledge of specific,
interactive behaviors and animal
physiology is known. Here are brief
descriptions of three studies being
conducted at or by the Saint Louis Zoo.
For more information about these or
other studies, visit at the Saint Louis Zoo
online at www.stlzoo.org.
ZOO
MOTHER / INFANT ANTELOPE
Saint Louis
101
ZOO STUDY
Saint Louis Zoo researchers are conducting studies at Washington University’s Tyson
Research Center to increase the numbers of the endangered Mexican wolves. Scientists
have a strong interest in understanding the behaviors related to reproduction
in order to support breeding efforts at the Wolf Sanctuary.
Researchers count the occurrence and duration of behaviors on a clipboard
and simple data sheet.
Previous zoo studies have focused on the
great Indian hornbill. The zoo is satisfied with
the results of this species’ breeding efforts.
More recently, the zoo began looking at
another hornbill species whose reproductive
habits are quite similar — the Blyth’s
hornbill. In fully understanding the intricate
behaviors and rituals of these birds, we can
better provide the environment for their
breeding success. We can also help other zoos
and in situ (in the country of origin)
conservation efforts find similar success.
The reproductive behaviors of hornbills are
fascinating. They are housed in mating
pairs, since these birds are among those
who will form monogamous pair bonds.
Females “interview” potential mates by mimicking the behaviors they will use during
nesting. A male will appropriately respond by exhibiting food-giving behavior, proving that
he will care for the female when she walls herself into a log/tree cavity, leaving only a
small opening through which she will receive food and carefully excrete her waste. During
this time she will lay her eggs, incubate and hatch them. She will emerge when the chicks
are ready to fledge.
Researchers collect data on a laptop computer.
CAREERS INanimal behavior
T hese careers require a doctor
of philosophy (Ph.D.) or doctor
of veterinary medicine (D.V.M.)
degree.
• Animal psychologist
• College professor/researcher in
biology, zoology, psychology,
others
• Conservation field researcher
• Health researcher
• Research assistant
• Veterinarian
• Zoo curator or researcher
• Zoo director
T hese careers require at least
a bachelor’s degree, and
probably a master’s degree.
• Animal care facility manager
• Animal caretaker
• Animal control officer
• Animal caretaker
• Companion animal trainer
• Veterinary assistant
• Zoo educator
• Zookeeper
from http://acunix.wheatonma.edu/kmorgan/AB_Careers/animal_behavior_careers.html.
MEXICAN WOLF
ZOO STUDY BLYTH’S HORNBILL
101
102
QA
What is an ETHOGRAM?
Ethograms are essential tools for
scientists and students who want
to study animal behavior and
communicate their results to others. A
complete ethogram describes the social
or environmental setting for each
behavior, includes information about
the timing and frequency of the
behaviors, and organizes the behaviors
into functional categories.
When we suggest a function for a
behavior, we are making an implicit
hypothesis that may require further
observation to verify. The name we
assign to a behavior can also imply a
function for that behavior. For example
the same body posture can be labeled
“lying down” and “resting.” “Lying down”
only describes what the animal is doing;
“resting” implies why the animal is doing
it. In fact many animals lie down for
reasons other than rest. Social animals
may lie down as a sign of submission
and predators or prey may lie down to
hide from each other.
In order to be a useful scientific tool for
studying behavior, an ethogram must be
unambiguous in distinguishing different
behaviors. Descriptions of behaviors
must be clear and complete. Data
shown in graphs and tables helps
others understand it better. Ethograms
can also contain pictures, or drawings
of the animal to help the reader
understand exactly what behavior is
being described.
AN ETHOGRAM IS A DESCRIPTIVELIST OF ALL BEHAVIORS OF APARTICULAR SPECIES.
habitat and basic behavior: While some monkeys can live in urban areas, the
lion-tailed macaque is a true rain-forest dweller and is unable to adapt to cities. All
macaques are primarily diurnal, and all have an arboreal capability, but most species
come down from the trees at least on occasion to forage or move over long distances.
The lion-tailed macaque spends less than one percent of its time on the ground. They
carefully inspect surroundings for a long time before descending from the trees.
Lion-tails have cheek pouches which they quickly stuff as they forage for food in
dangerous places. They are good swimmers. At night, they huddle in a sleeping
cluster high in the canopy. In captivity, the lion-tailed macaque has been observed
manufacturing and using tools to extract syrup from containers.
Macaques live in social groups of 10-20, including one to three adult males.
Females do not have their first offspring until they are 5 years old and males do not
reach sexual maturity until 8 years old. The species has a low reproductive rate,
which may contribute to its endangered status.
threats: Major threat is habitat loss to hydroelectric power, timber and tea
plantations, and harvesting of firewood and other forest products for human use.
They are also hunted by the Nilgiri hills natives for their skin and meat.
from Ali, Rauf. Lion-Tailed Macaque: Status and Conservation. 1985, Alan R. Liss, Inc. New YorkNowak, R.M. “Walker’s Mammals of the World," 6th ed. John Hopkins Univ. Press, Baltimore, 1999Singapore Zoological Gardens Docents, 2000, http://www.szdocent.org/pp/p-mclion.htm
lion-tailed macaqueMacaca silenus
107
appearance: Goat like mammals, with short, woolly undercoats and long coarse
guard hairs. Vary in color from gray to dark brown to reddish, with white or yellow throat
patches. Both sexes have conical horns, 127-78 mm in length, which curve toward the
rear. Stout, long limbs are well adapted to climbing.
size: Head and body 820-1200 mm, tail 76-203 mm, shoulder height 570-785 mm,
weight 22-35 kg
food: Diet consists of twigs, low shrubs, grass and nuts.
range: Gorals are found across most of the southern slopes of the Himalayas of
northern India, west China, and north to Korea, at elevations of 1,800-2,000 m.
habitat and basic behavior: Gorals tend to rest on sunny rock ledges,
where they are difficult to recognize even in full view. They are found on rugged,
wooded mountains, and seem to prefer the most difficult terrain possible. When
motionless, their color blends with the rocks. They live together in groups of
4-12, but males commonly live alone most of the year. When frightened, gorals emit a
hissing or sneezing sound.
They may spend most of the year within a home range of just a few dozen hectares,
but move a few kilometers to a more favorable wintering areas. They are most active
during early morning and late evening, but on cloudy days roam throughout the day.
After eating in the morning, they usually drink water and then retire to a sunny rock
ledge. They can be difficult to recognize in full view, because their color blends with
the rocks.
Gorals reach sexual maturity at age 3. Gestation lasts 6-8 months. One to two
offspring are born.
threats: Gorals are often hunted for meat and sport, even in protected areas. The
most significant threat to them is severe habitat disturbance and alteration, particularly
in the lower portions of the Himalayas and in northeastern India. However, goral can
survive in areas that have limited disturbance of shrubs and forest cover.
goralNaemorhedus goral
from Shackelton, D. M. (ed.) and the IUCN/SSC Caprinae Specialist Group. ”Wild Sheep and Goats and Their Relatives. Status Surveyand Conservation Action Plan for Caprinae." IUCN, Gland, Switzerland and Cambridge, UK, 1997. Nowak, R.M. “Walker’s Mammalsof the World," (6th ed.), John Hopkins University Press, Baltimore, 1999. Mochi, U. and T.D. Carter. “Hoofed Mammals of the World." Singapore Zoological Gardens Docents, 2000, http://www.szdocent.org/pp/p-mclion.htm
108
appearance: Monkeys with long
pointed muzzles, large eyes, and
triangular ears. Upper parts are brownish
gray, underparts are whitish, tail ringed
with black and white. Palms and soles are
long, smooth and leatherlike.
size: Head and body length 385-455
mm, tail length 560-624 mm, weight
2.3-3.5 kg.
food: fruits, leaves, other plant parts;
rarely insects. The kiley tree is the
mainstay of their diet.
range: southwestern Madagascar, in
wooded and thinly wooded country
habitat and basic behavior:Found only in Madagascar, lemurs are
among the most diverse primate faunas
on earth, and highly unusual. They live in
large family groups called troops,
consisting of 5-30 lemurs. The troops
forage for food throughout the day, and
can be active at night, as well.
Troops have no consistent leadership,
and feature much agonistic behavior and
fighting. The sexes have separate
dominance hierarchies, with females
dominating the males. The troops are
organized around a core group of adult
females and their infants. Females
remain in the troop of their birth, but
males move among the troops.
Lemurs have been considered as
territorial, however, territories overlap
considerably. Lemurs vigorously defend
areas being used. Disputes generally
involve two opposing groups of females
running at each other and vocalizing, but
direct contact is rare. Researchers have
identified 15 different vocalizations,
including a howl audible to humans at
1,000 meters. Mating occurs in spring
and births in late summer to fall. Births
often occur within a period of days in a
troop. Single young are most common,
but twins are not rare. Weight at birth is
50-80 grams. During the first two weeks of
life, the young cling to the mother’s
underside, then rides on her back.
Females generally conceive at 19-20
months. Males are sexually mature at age
2.5, but may not be allowed to mate by
older males.
threats: Madagascar is one of the
world’s most threatened biodiversity
“hotspots,” with 80 percent of its forests
already gone and most of what remains
at great risk. The ring-tailed lemur is the
only surviving semi-terrestrial diurnal
lemur. Its preferred habitats, of forests
along rivers, are disappearing because of
fires, overgrazing, and cutting trees for
charcoal. Hunting for pets and zoos is
also a threat.
from Mittermeier, R.A., I. Tattersall, W.R. Konstant, D.M. Meyers andR.B. Mast. Lemurs of Madagascar. Washington D.C., ConservationInternational, 1994.Nowak, R.M. “Walker’s Mammals of the World," (6th ed.), JohnHopkins University Press, Baltimore, 1999.
ring-tailed lemurLemur catta
109
butterflies and mothsMultiple species within order Lepidoptera
note: More than 160,000 species exist within Lepidoptera; students may compare
and contrast two or more species.
appearance: Insects with large, easily distinguishable wings. Among the most
popular and easily recognized insects, butterflies and moths are separated mostly by
observable differences. Most butterflies fly by day and most moths fly at night. Most
butterflies are brightly colored and most moths are dull. Despite these differences and
others, there is no single feature that separates all butterflies from all moths.
size: Range of sizes from very small to wingspans of 180 mm Idea tamb sisiana.
food: generally plants. The Pyralid (Laetilia coccidivora) is carnivorous; in the
caterpillar stage, it eats scale insects and aphids.
range: Butterflies and moths are found around the world.
habitat and basic behavior: Butterflies and moths each go through four
different life stages: egg, caterpillar, pupa, and adult. Life spans vary greatly between
species, from a few weeks or several years. Some of the wood-boring larvae of the
Cossid moths may spend months, or even years, in the caterpillar stage.
Many moths and butterflies have complicated courtship behavior, including
performing elaborate flights and “dances.” They often use chemicals called
pheremones to attract members of the opposite sex. When a male finds a female
who shows interest in him, they both land. The mating pair will often tap each other
with their antennae, detecting other scents. Mating may last for 20 minutes to
several hours, during which time the insects do not move. Most males proceed to
look for other females after mating, but females seek a place to lay their eggs.
Some scatter their eggs, but others seek a food plant for the new caterpillars.
Some butterflies, such as the Orangetip (Anthocharis cardamines), have distinctly
different appearances for males and female. This is known as sexual dimorphism.
Whalley, P. Butterfly and moth. New York: Dorling Kindersley, 2000.Feltwell, J. The encyclopedia of butterflies. Great Britain: Quarto Publishing, 1993.
110
appearance: Mammal with prominent whiskers, medium size eyes, small ears, thick neck and long,
heavy tail. Legs are short with webbed feet. Color is dark brown or black, light brown or gray on belly, and
silvery on face or throat.
size: 90-135 cm with one-third of length consisting of the tail; 4.5-11.5 kg
food: Mostly fish and crayfish, sometimes frogs, salamanders, snails, clams, snakes, turtles, muskrats,
birds, insects, earthworms
range: North American lakes and rivers
habitat and basic behavior: Otters usually live in family groups. They live in burrows in the
banks of rivers or lakes, usually ones that were built earlier by muskrats, beavers or woodchucks. Home
ranges may include 80-160 miles of shoreline. Otters are powerful swimmers that often swim with just the
top of the head and eyes above water. They can swim 10 km/hour on the surface and submerge for up
to 4 minutes.
Even the adults regularly engage in social behaviors that are best described as play. They are particularly
fond of sliding down steep slopes that have been made slippery with mud or ice, and that terminate either
in a deep pool of water or snowdrift. They will climb up such a slope and slide back down repeatedly,
particularly if accompanied by other members of the group. They also spend prolonged periods in small
groups tossing rocks or clam shells into the water and then diving for them. One of the most common ways
of traveling on snow is to run for a few feet, then slide on their bellies as far as they can.
threats: Hunting and trapping for fur has greatly reduced populations, and river otters are considered
endangered today. They are parasitized by tapeworms.
Saint Louis Zoo
river otterLutra canadensis
111
appearance: Cylindrical mammal
completely lacking hair, except for a few
sensory whiskers (vibrissae) on face and
tail, plus a fringe of hair on feet. Short
legs, tiny eyes and no external ears. Very
powerful jaw muscles and large
protruding incisors used for burrowing.
Skin is very wrinkled because no fat layer
is underneath. Color varies from pink to
yellow to white or gray.
size: 7.0-8.5 cm body length, tail length
3.5-4.0 cm, weight 30-60 gm; queen
may be 90 gm.
food: Mole-rats are strict herbivores;
food includes roots, tubers and
underground plant parts found while
burrowing.
range: Eastern sub-Saharan Africa,
central and eastern Ethiopia, central
Somalia and Kenya.
habitat and basic behavior:Naked mole-rats are the only mammals
that exhibit eusocial behavior. This type
of behavior describes the colony
lifestyles of bees, ants and termites.
Eusocial animals live in large colonies
within which only one female, the
queen, and a small number of males
participate in reproduction. The rest of
the colony members share such tasks
as feeding and defending the queen
and raising her offspring. In honey bee
colonies, the queen produces a
chemical which prevents other females
from breeding. In naked mole-rat
colonies, the queen fights other
females, often to the death, to prevent
them from breeding.
A queen may breed for 12 years, or until
she is displaced by a more aggressive
female. The queen has four litters a
year, with an average size of seven to
ten pups. When a litter is born, a group
of non-reproductive colony members
forms a living carpet on which the
queen and her pups lie for three to four
weeks while the queen nurses them.
The first solid food the pups eat are
feces that they actively solicit the adults
to produce for them. The young then
begin to help keep the tunnels clear and
to gather food for the nest. Older
individuals dig new tunnels in search of
food and defend the burrow against
intruders.
threats: Burrowing reptiles,
especially the rufous-beaked snake.
Saint Louis Zoo
naked mole ratHeterocephalus glaber
112
goatEight species in genus Capra
appearance: Goats with varied sizes and horns. Males
are odorous, with beards. In wild species, horns of males are
500-1,600 mm long and horns of females are 150-380 mm
long, with much variation between species. Some are long
and curved, some curlicued.
size: Sizes are varied, but males are generally larger than
females.
food: All species graze for grass and vegetation.
range: Europe, Asia, and Africa
habitat and basic behavior: Most goats are well
adapted to mountainous areas and are good climbers. Adult
males and females are often separate for most of the year.
The Cretan wild goat (Capra aegagrus) lives in groups of five
to 25 individuals, which vary through the year. Females
remain in the groups except to give birth. Males usually stay
apart in small bachelor groups. They form dominance
hierarchies and compete for mating privileges. Conflicts
involve threats, visual displays with the horns. They may rear
up and crash horns together.
threats: Hunting for horn trophies and meat has
decimated many species. The domestic goat is maintained
worldwide for milk, meat and wool. It often competes with
wild species for food. The European ibex was reduced by
sport hunting and a demand for body parts to only about 60
individuals in northern Italy in the nineteenth century. Those
animals were carefully protected and used to reintroduce
the species to the Alps. By the 1980s there were more than
12,000 in Switzerland and elsewhere.
from Nowak, R.M. “Walker’s Mammals of the World," (6th ed.), John Hopkins University Press,Baltimore, 1999.
113
chimpanzeePan troglodytes
appearance: Monkeys with
prominent ears, protruding lips, arms
that are longer than legs, a long hand
with a short thumb, and no tails. Face
is bare and usually black, color ranges
from deep black to light brown. Hair
on the head may grow in any
direction, and baldness occurs in both
sexes at maturity.
size: Head and body length about
635-940 mm, height 1.0-1.7 meters,
weight 34-70 kg for males, 26-50 kg
for females. Captive animals can
weigh as much as 68-80 kg.
food: Fruit, seeds, honey, insects,
eggs and meat. They may prey on
other monkeys and small animals.
range: Tropical rainforests and
savannah in Gambia and Uganda
habitat and basic behavior:Most chimpanzees spend daylight
hours in trees. Young individuals may
swing from branch to branch
(brachiate), but the most common
form of locomotion is knuckle-walking:
walking on all fours with the fingers
turned under. They live in highly
socialized societies that may have as
many as 80 members. Social position
in the hierarchy is extremely important.
Chimps communicate by facial
expressions, vocal sounds and posture.
They have a long childhood, with
young dependent on their mothers
until age 5, and maturity at age 13.
Bonds between mother and young,
especially females, can last a lifetime.
Chimps can live up to age 50.
The chimpanzee is renowned for its
ability to make and use tools. They often
feed by carefully poking a stick or vine
into a termite nest to collect the insects.
Chimps also use sticks as hooks to pull
down fruit-laden branches. They may
fight with sticks as weapons. They use
stones as hammers to crack nuts.
threats: Hunting for pets, and
habitat loss threaten the chimpanzee in
the wild.
from Nowak, R.M. “Walker’s Mammals of the World," (6thed.), John Hopkins University Press, Baltimore, 1999.
114
malayan sun bearUrsus malayanus
appearance: These small bears are mostly
black, with whitish or orange breast mark and
grayish or orange muzzle. The body is stocky,
muzzle short, paws large and claws are strongly
curved and pointed. The soles are naked.
size: 1,000-1,400 mm, tail length 30-70 mm,
shoulder height about 700 mm, weight 27-65 kg.
food: omnivorous, including honey, insects, larvae,
coconut palm, termites, fowl, rodents, fruit juices
range: Dense forests in central China, Burma,
Thailand, Indochina, Malay peninsula, Sumatra,
Borneo
habitat and basic behavior: Sun bears
are active at night, usually sleeping and sunbathing
by day in trees. They break or bend tree branches
to form nests and lookout posts. The walking gait is
unusual in that all feet are turned inward. The
species is very shy and retiring and does not
hibernate. An expert tree climber, it is cautious,
wary and intelligent. A young captive observed the
way in which a cupboard containing sugar was
locked with a key, and later opened the cupboard
by inserting a claw in the keyhole and turning.
Mothers give birth to one or two young, about
325 gm each. Cubs are blind, nearly naked and
helpless. They are very dependent on their
mothers for at least three or four weeks. Once out
of the den, cubs learn to climb very quickly. They
stay with their mothers for at least a year, possibly
longer. Captive bears can live up to 31 years
threats: Sun bear cubs are popular as pets, but
larger animals are unruly and often sold as parts,
which are used in the medicinal trade. Logging of
forests and conversion to rubber plants has
contributed to habitat loss and population decline.
They are considered endangered. Tigers and other
big cats are predators.
from Nowak, R.M. “Walker’s Mammals of the World," (6th ed.), John HopkinsUniversity Press, Baltimore, 1999.Stonehouse, B. Bears: A Visual Introduction to Bears. AND Cartographic Publishers Ltd. New York, 1998.
115
appearance: Large, brilliantly colored aquatic birds. Long sinuous necks, long
legs and webbed feet. Slim, rose colored wader. Bill is specialized for filter feeding
and sharply bent in the middle.
size: 90-155 cm tall, males are a little larger than females.
food: Small marine life, including mollusks, crustaceans, blue-green algae and
diatoms.
range: Mudflat areas of southern Florida, South America, Africa and Asia. Highly
adaptive to chemicals in water, including chlorides, sodium carbonate and sulfides.
Can exist in very hot and high altitude climates.
habitat and basic behavior: Family is one of the oldest bird groups still
alive, with fossil evidence going back to early tertiary. The birds are highly social and
gregarious, living in large flocks of up to tens of thousands. In east Africa, over a million
lesser flamingoes can gather. Birds often spend the day resting, and feed at night.
Starting several months before breeding and continuing afterwards, flamingoes often
devote considerable time to the performance of collective displays, which may be
carried out by hundreds of birds. They consist of a series of ritualized postures and
movements. One of the most common, “head flagging,” involves stretching the neck
and head up as high as possible and turning head from side to side. The purpose of
these displays is to synchronize, by means of hormonal stimulation, the breeding
attempts of as many birds as possible. In this way, populations can make the most
of the periods when conditions are optimal.
A characteristic feature of flamingoes is their resting posture of standing on one leg.
In this way, the amount of heat lost through the leg and foot is reduced. This
posture is also used in hot weather, so it is likely comfortable for the birds.
threats: Humans have been mostly threatened by human capture for zoos.
Wetland habitat destruction also threatens the birds.
from Harrison, Bird Families of the World, New York: Abrams, 1978.del Hoyo, J., Elliott, A & Sargatal, J. eds. Handbook of the Birds of the World, Vol. 1. Lynx Edcions, Barcelona, 1992.