September 2001

BIO 3136 - ANIMAL BEHAVIOR (Course outline) Instructor: Dr. Jaroslav Picman

Vanier Rm. 305, 307

Tel: 562-5800 (4548) E-mail: [email protected]

Demonstrator: Sarah Derrane

Vanier Rm. 313

Tel: 562-5800 (4575) E-mail: [email protected]

Class times: Lecture: Lecture/Film: Discussions:

Tue Fri Mon Mon Tue Wed

8:30-10:00 10:00-11:30 10:00-11:00 16:00-17:00 11:30-12:30 13:00-14:00

MRT 219 MRT 219 CBY B202 CBY E016 MCD 120 LMX 112

Text: John Alcock. 2001. Animal Behavior; An Evolutionary Approach. Seventh edition. Sinauer Associates, Inc. [Note: Purchase a copy of this text immediately, if you have not done so already. You will be asked to read several chapters each week, starting the first week.] Course content: The course aims to introduce you to the study of animal behavior by offering you a variety of learning situations. The course will consist of: [a] lectures; [b] an independent research project describing the behavior of an animal of your choice; [c] discussions based on selected material from the text and original research articles; and [d] films. The individual aspects of the course are outlined in more detail below. a] Lectures: There will be one to two lectures per week, regularly each Tuesday and every second Friday (see above for timing and location of our meetings). The aim of these lectures will be to deal with Page 1

selected topics in depth to show how behavior studies are carried out. Although the lectures will not aim for complete coverage, they will be synthetic as far as possible, stressing generalities. You can purchase a copy of lecture presentations for BIO 3136 from Burney (Xeroxing facility in Vachon Bldg.). b] Project: (see separate information on project!) Most accounts of what animals do appear deceptively clear and straightforward. In practice, however, how one interprets an animal’s behavior depends on how one observes and records it. The only effective way for you to understand the complexity of the problem involved is for you to try it yourself. Because space limitations prevent a formal lab, you will carry out a brief project describing some aspect of the behavior of an animal of your choice and, hopefully, testing some of your ideas on the function/adaptive value of that behavior. Towards the end of the term, you will be asked to give a brief (around 15 min) verbal report on your project and submit a written report. c] Directed study from the text: (see the study guide to the text) The whole class will be divided into three to four groups (in each group a maximum of 12 students). These groups will be organized during the first week of classes. The group members will meet with their instructor once a week. The discussion groups are designed to serve as a forum for the following activities: 1) Directed study from the text; During weeks 1-7 you will be asked to read certain sections of the textbook. During the discussion group meetings we will discuss material covered and any other problems arising from the text. The purpose of this exercise is to give you a brief overview of the subject and familiarize you with the necessary jargon of behavior studies. Although you will be asked to do an extensive amount of reading during the first seven weeks, this is absolutely essential because the information that you will master is going to prepare you for your research projects. Fortunately, the text is excellent and extremely interesting! 2) Discussion and supervision of project activities; Your discussion group leader will also be responsible for giving you project advice, and helping you with solving problems, whenever these arise. The verbal presentation of your planned research activities (a research proposal) and the final report on your findings will be presented in front of your discussion group. Your group leader will evaluate the written report on your project. 3) Discussion of selected topics; After the completion of the text study, the discussion groups will be used as a forum for discussions of several topics, selected original research articles (critical evaluation of two selected original research articles with the purpose to demonstrate how research papers in the field of Animal Behavior are organized and written), and for debates and practice in writing essays and project reports. d] Films: Page 2

Films are an extremely useful aid particularly in this course. Films selected for the course will effectively demonstrate various behavioral patterns and their function in lives of various animals. Several films will be shown on selected Thursdays, starting the third week of school. If we were unable to obtain a particular film for a given meeting, we would let you know in advance about it. In the case of cancellation of a film presentation, we would use the meeting time for alternative activities such as a special lecture or discussion of research projects. The instructor will remind you on Tuesday (lecture meeting) if any film will be shown next Friday. NOTICE: I recommend that you regularly attend the film sessions not only because selected films are excellent and very interesting, but also because the material covered in these films could be used in the second exam. Exam: There will be no scheduled final exam, instead there are going to be two class exams. One exam will be based on the study text only (first midterm) and the other on lecture material as well as on the study text (second midterm at the end of term). Because there is going to be no final exam, it is really important that you write both class exams. More details on these exams will be given later. [NOTE: Please, be aware of Faculty Regulations regarding attendance and examinations; see the appropriate sections in your 2001/2002 calendar. Furthermore, with respect to students’ requests for mark changes, please, be aware that the Faculty of Science has accepted a policy that, upon the instructor’s verification of an exam, the initial mark that was assigned can be either increased, can remain the same, or can be decreased.] Marks: Marks will be allocated as follows: 2 exams .............................. 50% (Each exam 25%) Project (written report) ....... 40% Discussion contribution ...... 10% (attendance 5%, discussion contribution 5%)

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A STUDY GUIDE TO ALCOCK’S TEXT Every student knows that lectures are an inefficient way of conveying factual information. One cannot listen and take notes effectively at the same time. On the other hand, to master any subject requires a baseline knowledge of facts, terms, and concepts. Since most of these things are available in the textbooks, it makes sense for you to acquire this information about behavior from a text, but to have an opportunity to discuss the material and apply it to simple problems. Alcock’s book is an up-to-date book which is well illustrated, referenced, and has a more synthetic treatment than most other texts. The evolutionary viewpoint taken by Alcock contrasts with more physiological ones represented in some other textbooks. The Alcock’s book is one of the best textbooks that is being used by many universities across North America. We will work through certain sections in Alcock on the following schedule. After each chapter block, there are the key concepts that you should attempt to grasp. Read for week No.

Chapters

Key concepts and terms

2

1,2,3

How and Why questions: proximate and ultimate causes of behavior; Darwinian argument on evolution; Individual selection; group selection; testing alternative hypotheses; certainty in science. Function of song in birds; song development; the role of physiology and environment in development of behavior; critical period; the role of genes in development of behavior; dialects. Genetics and behavior; genotype; phenotype; artificial selection.

3

4,5,6

Environment and the development of behavior; the interactive theory of development; the role of hormones in the development of behavior; imprinting; early learning; behavioral flexibility in predictable and unpredictable environment; developmental homeostasis. Nervous system and the control of behavior; FAP; stimulus; releaser; IRM; supernormal stimulus; neurons; action potential; synapse; stimulus filtering; selective visual perception; optical illusions; face detectors; navigation; orientation mechanisms. Organizing mechanisms controlling behavior; command centers; circadian mechanism; long-term cycles; the role of hormones.

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4

7,8,9

Adaptation; comparative method; cumulative selection; convergent and divergent evolution; direct and indirect measures of fitness; antipredator behavior; cryptic behavior; anti-detection behavior; warning coloration; Batesian mimicry; unprofitable prey model; vigilance; selfish herd effect; dilution effect; geometric effect; rapid escape; misdirecting attackers; game theory. Feeding behavior; locating food; honeybee dances; information center hypothesis; food selection; optimal foraging theory; optimality theory; habitat selection; ideal free, ideal despotic distribution; habitat preferences; dispersal mechanisms; inbreeding depression; migration; conditional strategy; territoriality; home range; ESS; the payoff asymmetry hypothesis.

5

10,11

The evolution of communication; dialects; adaptation; adaptationist approach;honest signals; deceptive signals. The evolution of reproductive behavior; why two sexes?; operational sex ratio; parental investment; sexual selection; alternative mating tactics; sperm competition; mate guarding; female choice; conflict betwen sexes.

6

12,13

Mating systems; monogamy, polygamy, polygyny, polyandry promiscuity; extra-pair copulations; sex role reversal; female defense polygyny; resource defense polygyny; scramble competition polygyny; lek polygyny. The two sexes and parental care; offspring recognition; brood parasitism; parental favoritism.

7

14

Benefits and costs of social behavior; helping behavior; altruism; indirect and direct selection; coefficient of relatedness; direct, indirect and inclusive fitness; helping; alarm calls; eusociality.

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BIO 3136 - GENERAL READING LIST BOOKS: Alcock, J. 1998. Animal Behaviour, An Evolutionary Approach. Sinauer. Sixth edition. The study text. Barash, D.P. 1962. Sociobiology and Behavior. Elsevier. An excellent, ecological1y oriented book. Brown, L. and J.F. Downhower. 1988. Analyses in Behavioral Ecology. A manual for lab and field. Sinauer Associates, Inc. Brown, J.L. 1975. The Evolution of Behavior. Norton. Deals more extensively with ecological aspects of behavior. Dawkins, R. 1976. The Selfish Gene. Oxford. A popular book promoting the idea that natural selection acts on individual genes. Excellent - highly recommended! Drickamer, L.C., Vessey, S.H., and E.M. Jacob. 2002. Animal Behavior; Mechanisms, Ecology, Evolution. Fifth edition. McGraw-Hill. A general textbook. Gould, J.L. 1982. Ethology; The Mechanisms and Evolution of Behavior. W.W. Norton & Company. Stressing evolutionary aspects of behavior. Goodenough, J., McGutee, R., and R. Wallace. 2001. Perspectives on Animal Behavior. Second edition. Wiley. Grier, J.W., and T. Burk. 1992. Biology of Animal Behavior. Mosby Year Book. Krebs, J.R. and N.B. Davies. 1978. Behavioural Ecology; An Evolutionary Approach. Ecologically oriented, more advanced. Krebs, J.R. and N.B. Davies. 1993. An Introduction to Behavioural Ecology. Sinauer. An excellent book on Ecology of Animal Behavior. McFarland, D. 1999. Animal Behavior. Pitnan. 3rd edition. Generally an excellent textbook. Wilson, E.O. 1975. Sociobiology, the new synthesis. Harvard. An excellent survey of social behavior in animals. Wittenberger, J.F. 1981. Animal Social Behavior. Duxbury Press. An exhaustive account of all major hypotheses; highly advanced; excellent. JOURNALS: Most original research papers could be found in the following specialized journals: Animal Behaviour, Behaviour, Behavioral Ecology, Behavioural Ecology and Sociobiology. Many papers dealing with behavioral research may also be found in more general journals such as The Auk, Canadian Journal of Zoology, Condor, Ecology, Oikos, Wilson Bulletin, etc. I strongly recommend that you spend some time going through the recent issues of these journals to find out what is being done in this field of biology. This exercise is also likely to help you with finding a suitable topic for your research project. TV: There are many excellent movies shown on TV that deal with various aspects of animal behavior and ecology. Examine the TV guide carefully for these programs. Page 6

SUMMARY OF LECTURE MATERIAL The following is a tentative list of lectures to be given in BIO 3136: Date

Week

Lecture topic

Sept 7

1

General introduction; Organizing discussion groups

11 14

2

History of Animal Behavior; How and Why questions Adaptation; Selection theory; Genetic basis of behavior; Heritability concept

18 21

3

Spacing systems I. MOVIE SESSION 1

25 28

4

Spacing systems II. - Territoriality Habitat selection

2 5

5

Grouping behavior MOVIE SESSION 2

9 12

6

Sexual reproduction Mating systems I

16 19

7

Mating systems II Univ. of Ottawa Day - CLASSES CANCELLED

23 26

8

Alternative mating strategies MOVIE SESSION 3

30 Nov 2

9

E X A M I (Based on textbook only) Research Projects - How to prepare reports; Data analyzes

6 9

10

Sexual dimorphism MOVIE SESSION 4

13 16

11

Brood Parasitism - an alternative reproductive strategy Ecology, Behavior and Conservation Biology - Why are some of our birds disappearing?

20 23

12

Review session - preparation for exam II MOVIE SESSION 5

27 30

13

E X A M II (based on text and lecture material) Research projects

Oct

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BIO 3136 - PROJECT For practical reasons it is not possible to arrange a formal laboratory for the Animal Behavior class. Nevertheless, it is important for students taking the course to have some preliminary experience of the observation, description, and recording of behavior. The following is an outline of a project to be carried out over the next few months and completed before the end of term. OBJECTIVE: Observe, describe, and record the behavior of an animal of your choice. Suggested subjects: beaver; raccoon; hamster; mouse; cage birds; aquarium fish; wild birds such as ducks, swans, gulls, herons, pigeons, house sparrows, chickadees; birds at a feeding station close to a window; farm animals; or any animal that you have convenient access to. Although cats and dogs are suitable subjects, in general we will not encourage students to conduct their projects on these household pets because: (1) people find it difficult to be objective in describing their own dog or cat; and (2) observations are restricted to a single individual whose behavior is so obviously conditioned to its local environment. OUTLINE: In most cases work should fall into two phases: (1) Observation and description (2) An attempt to take a quantitative record. This may simply be to convince others of the accuracy of your description, or may involve a simple experiment designed to answer some question(s) suggested by your preliminary observations. We do not expect a detailed comprehensive study of a whole species' repertoire. It will probably be more fruitful for you to first make a few general observations on your chosen animal and then pick a particular behavior for a more careful study. EXAMPLE: Observe the behavior of a hamster or gerbil, look for patterns of activity during the day, observe the behavior patterns involved in feeding, washing, exploratory behavior, reactions to other animals, etc. You might then concentrate on feeding behavior - the motor patterns involved, food selection, etc. This might suggest simple experiments on which foods your animal(s) prefers in a choice situation, or on the way it recognizes food, etc. Page 8

It is very important that we discuss your project with you. Before you start collecting data, your project must be approved by the leader of your discussion group. PROJECT ADVISING should normally be done by the leader of your discussion section. Some class time will be spent on this topic during term. NOTE: All projects involving animals must first be approved by the Animal Care Committee. For this reason, we will need a short description of the key objectives and methods to be used in your study (1-2 pages of text; 2 copies), that we will have to submit for approval to the local A.C.C. We will devote some class time to this problem and prepare you for this activity to increase the chances that your project will be approved. REFERENCES: Preferably none. The purpose of this exercise is not to conduct an original piece of research but to expose you to a new learning experience. Examples in the text or other reading material should lead you to many potential topics. In general, we would prefer that you do not start by trying to read research papers about your chosen animal. One reason for this is that this would bias you to look for what others have seen so that you overlook something that is really happening but what the previous observers missed. Naturally, we hope that you will be sufficiently interested to want to look at literature dealing with your problem at a later stage in your investigation. You may find, however, that there are no published accounts. The final result will be evaluated on what you have done, rather than what you have read of the background material. ANIMALS AND EQUIPMENT: No animals or animal facilities will be provided by the Biology Department. Therefore, all projects will have to involve wild animals, or, if you have an access to farm animals or suitable pets, then these could also be used (but only after approval by your instructor). In general, we will not approve any project during which the animal's welfare would be adversely affected (e.g. studies involving starvation, or studies that would physically restrict animals in any way). Details will be given in class by your Instructor. NOTES ON PROCEDURES: Once you have familiarized yourself with the major activities of the animals you are studying, write down detailed descriptions of the behavioral patterns in such a way that someone else reading your account will be able to recognize the events you described. The written description should be as concise as possible and will almost certainly be helped by drawings or simple diagrams. Page 9

NAMES: Name the behavioral patterns you have described on the basis of what you see, rather than on some interpretation of what you think it means. E.g. if a cat bares its teeth on meeting another cat, call it "bare-toothed posture' and not "aggressive threat". RECORDING PROCEDURES: You should quantify your observations for two reasons: (1) This will improve the accuracy of your description. Rather than saying that a fish sometimes swims, you can express this as a percentage of the total observation time. A1ternative1y, you may want to ask whether there are particular sequences of actions which tend to go together. To answer this, you must count how often behavior A follows behavior B, etc. (2) It is almost impossible to answer any experimental question without a quantitative measure of the effects of your treatments or details of your measures. RECORDING METHODS: (1) After a preliminary study of your animal, develop a shorthand or a series of symbols to Identify the behavior patterns you see. (2) Take a continuous record for a period long enough to give you an adequate sample of the actions you wish to quantify. This will obviously vary from case to case; you might describe the of a fly adequately on the basis of 10 minute observation, but it would take years to describe the social behavior of elephants. If behavior happens very fast, you may have to treat it as a series of point events which you can then sample at regular (e.g. 5 second) intervals. A time base is usually helpful and pre-prepared data sheet and a watch with a second hand may be helpful in providing this. (3) From your continuous record, you may extract the following kinds of information: (a) Frequencies: number of times certain events occur per unit Page 10

of time; absolute or relative frequency. (b) Sequence:

analysis of the temporal order of events.

(c) Temporal pattern: similar to sequence, but incorporating details of the durations of events and the time intervals separating elements of a sequence. Which of these you choose depends upon your animal and on objectives of your project.

TIME COMMITMENT: We expect about two weekends' (a minimum of one weekend!) worth of data collection, or an equivalent amount of time spread over more days. Plan your observations carefully, and do not fall into the trap of collecting more information than you have time to analyze. Under certain circumstances (e.g. in case of closely related projects) we may allow two students to cooperate in designing their project and collecting data. Data analyzes, interpretation of results and preparation of the final verbal and written report will, however, be done by each individual separately. VERBAL REPORT: During the last two to three weeks, the discussion groups will meet to give you an opportunity of telling others about your project. Each person will talk for about 10 minutes on some aspect of his/her observations. The 10-minute talk will be followed by a 5-minute discussion session. This presentation should help you to improve the final written report. WRITTEN REPORT: The written report must not exceed 3,000 words (i.e. 10 double-spaced pages of text). Long reports will loose marks, no matter how good they are. If you are having trouble keeping it down, leave some of it out. Each report must have the following sections: ABSTRACT, INTRODUCTION, METHODS, RESULTS, DISCUSSION (or a combined section RESULTS AND DISCUSSION), and REFERENCES CITED. If it helped the reader, then further subdivide the main sections into subsections (use headings that describe well the material treated in subsections). Also, when citing data obtained by other researchers, do not forget to state the source of information in a conventional way (name of the author and year). A piece of advice: use one of the original research articles as an example of how to Page 11

organize and write a scientific paper!!! Use tables and figures (diagrams) to give a clear and economical presentation of information. All Tables and Figures must be numbered, must have descriptive heading, must be well explained, and all must be referred to in the text (i.e. in the RESULTS). Again, have a look at your 'favorite" original scientific article published recently and use it as an example when preparing Figures and Tables. If possible, type your report and keep a copy for yourself (we will keep the original copy with class records, but you will be given an evaluation sheet). WHEN TO START? Do not delay too long in selecting your study species. Begin with observations as soon as possible. The timing of data collection will obviously be determined by nature of your project. The written report must be submitted during or before the last scheduled class meeting to your group instructor, who will be responsible for its evaluation.

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ANIMAL BEHAVIOR (BIO 3136) SUGGESTIONS FOR PROJECTS (1) (2) (3) (4) (5) (6) (7) (8) (9)

Food preferences in ................ Effect of weather conditions on feeding behavior of ............. Effect of time of day on foraging activities of ........... Hoarding behavior in squirrels. Spacing behavior in roosting pigeons, etc. ........... Aggressive behavior in mixed-species flocks at a feeding station. Effects of interspecific interactions on the spacing behavior of .......... Habitat preferences by .......... Feeding behavior of chickadees (you could also study any of the local waterfowl species); effect of interspecific interactions; effect of group size; effect of risks of predation, etc. ............... (10) Effect of flock size and food availability on foraging behavior of gulls, pigeons, sparrows, etc. (11) Foraging behavior of beavers.(based on signs of foraging activities around a beaver colony). (12) Adaptive value of spacing behavior in ......... (13) Roosting behavior in pigeons. (14) Local movement and competition of house sparrow flocks. (15) Foraging activities of swans. Behavioral interactions between male and female mute swans; effect of ... (presence of competitors, food provisioning, etc) on ...... (16) Foraging behavior of honeybees. (17) The effect of danger of predation on foraging activity of chickadees on a feeding station. (18) Exploratory behavior of mice, hamsters, fish, etc. (19) Factors influencing territory selection in ..... (some strongly territorial fish species, if you happen to have some at home). (20) Home range and competition of house sparrow flocks. (21) The effect of habitat complexity and type of release on density of territorial three-spined sticklebacks. (22) Effect of habitat complexity on spacing behavior of ..... (23) Learning in .......... (24) Factors affecting coloration changes in Anolis lizards. (25) Webb building behavior of spiders. (26) Dominance hierarchy in .... (horses, cows, sheep, chicken, etc......). (27) The effect of pheromons on spatial activities of ..... (mice, gerbils, etc.). (28) Conflict behavior in .......(e.g. gulls foraging in an area with a predator) (29) Preening behavior of (parrots, etc.)..... ________________________________________________________________________ Page 13

NOTES: (1) The above list is by no means exhaustive. If you have some other idea(s) (this will definitely happen after you will have read several chapters of your textbook), you will be expected to meet with the instructor who will discuss these ideas with you and then make a decision whether or not a given idea is suitable for a project. A suitable idea is one that is scientifically sound and that is feasible (i.e. you must be able to collect enough data within a relatively short time period reserved for this exercise). (2) Try to find a topic for your project that would allow you to conduct a simple experiment. The experimental approach will make it possible for you to answer some of your questions directly (i.e. in a much more effective way). (3) No experiments involving starvation or other experimental manipulations that might endanger animals in any way will be allowed. (4) Select a topic that will allow you to study animals in their natural environment. Consider "laboratory" type of projects (e.g. those that would involve your caged pet or aquarium fish) only if you failed to find a suitable field study. (5) Start thinking about potential topics for your project immediately. The examination of original research papers published in journals such as Animal Behavior will also be extremely rewarding.

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