Animal Behavior - Biology 261

Instructor: Robert A. Mauck




Ethology, the comparative study of behavior, is a young science, and we may trace the genesis of modern ethology to Konrad Z. Lorenz.  Through seminal thinking and prolific writing, Dr. Lorenz and Dr. Nikolaas Tinbergen, Lorenz’s most influential student, have spread the concepts of ethology worldwide from their cradle in central Europe.

Ethologists are behavioral zoologists.  They seek the biological roots and meaning of what animals do.  Not surprisingly then, they feel that any experimental manipulation of behavior of a species should be performed only once that species’ natural behavioral repertoire is known intimately.  That is, a catalogue of 'normal' action patterns is necessary to provide controls against which manipulated behavior can be viewed.

Ethologists constructing ethograms are, almost by definition, patient.  Such an attribute is necessary to accomplish the task of observing and recording everything possible about the behavior of an animal species under natural or nearly natural conditions.  Many examples abound of the patience required.  One man for months on end broiled in a seaweed-covered blind under the desert sun of the Peruvian coast, downwind from heavily aromatic carcasses of horses (Equus caballus).  All this to record, during brief encounters, the behavior of a magnificent vulture, the Andean condor (Vultur gryphus), attracted to the horseflesh.

The product of observation, note-taking, and synthesis of behavior has been called an ethogram.  This term describes an annotated catalogue or list of behavior patterns condensed from field notes, grouped in some coherent fashion, and sufficient to characterize what a given species in a given environment does.  For this course, you will compile, edit, and produce an ethogram depicting the behavior of a single animal species.



Animal Subjects


Alone and in consultation with me, you should weigh several factors before choosing the subject species for your ethogram.  Variation in behavior among individuals is the rule, can be expected, and is important.  You will want the opportunity to observe several or more animals of the same species.  So long as your presence does not change the species’ behavior, you should try to observe at the closest range possible (and always downwind from mammals), so as to pick up the smallest details of movement.  If you need them, the department can furnish you with distance-decreasing-technology (e.g., binoculars).  If you don’t want to use such tools, be sure to select an animal species and situation allowing close approach.  For instance, undertaking an ethogram of the wary whitetail deer (Odocoileus virginianus) without binoculars would not seem to be a very good idea.

Ethologists believe that animal behavior is responsive to environmental selective forces.  Humans (Homo sapiens) have acted as a particularly strong environmental force in shaping the behavior of their domestic animals.  Compare, for instance, the temperaments of various breeds of dog (Canis domesticus).  I would prefer you to study non-domesticated species, but if this is impractical, keep clearly in mind the probability that behavior of domestics has been molded through time by humans.

Practicality dictates that within the time span allotted you should select a species you can count on to be available when you are able to observe it. For example, in the winter months large temperate-zone cities in the U.S. will provide for study many house sparrows (Passer domesticus) and few American robins (Turdus migratorius).

The disruption of normal behavior by the observer's presence must be guarded against at all costs.  In the hierarchy of species-environment pairs listed below, I urge you to choose as near the first alternative as possible.  I strongly discourage you from choosing a subject from categories four or five.


1.              ‘Wild’ animals in their natural environments.  View environmental quality from the perspective of the potential study species.  Times Square and 42nd Street is a perfectly natural environment for a pigeon (Columba livia).


2.              ‘Wild’ animals in large enclosures.  The notion of large is relative.  A 100-liter aquarium may be large for an adult crayfish (Cambarus sp.), but not for an adult painted turtle (Chrysemvq picta).


3.         Domestic animals free-ranging (e.g., dogs, chickens, Gallus gallus) or in large enclosures (e.g., horses, cattle, Bos taurus).  You should use good judgment here, particularly in considering whether your presence influences behavior.  A beagle dog ranging after rabbit (Sylvilagus floridanus) scent in a wood is a different animal behaviorally from a beagle ranging after a pair of slippers in your closet.  In the higher species (e.g., dogs, horses, crows, Corvus brachrhynchos) that interact strongly with humans, avoid studying your own animals.


4.              Domestic animals in small enclosures.  More desirable than #5, as many species have been selected partially for the ability to function while closely confined.


5.         Wild animals in small enclosures.  Although this is the situation often encountered in zoos, try to avoid it as such animals often develop atypical and sometimes even neurotic behavioral patterns.


For this class, you may not use humans as a study species.  Humans’ behavioral variability is too great to be encompassed adequately in a short study.  Also, the problem of the observer's presence influencing behavior is quite severe.



Equipment and Note-taking


An accurate, comprehensive ethogram depends upon keen observation and accurate, efficient notation.  Tape recorders are excellent for capturing information rapidly, but they have several considerable drawbacks.  They tend to malfunction in the rain!  On most models, the tape can run out without warning.  Unless you have a full-time secretary, transcribing notes from the tapes can be a colossal annoyance.  I suggest you take notes by hand directly into a notebook with sewn-in or glued-in pages.  Loose-leaf notebooks tend to lose pages in a strong wind.  I recommend you use a ballpoint pen with waterproof ink and a clear plastic shaft so you can monitor the ink supply.  A 'BIC' pen is excellent, but you may find it necessary in sub-freezing weather to warm it up periodically inside your glove.  A pencil is also adequate, but remove the eraser.  Erasing field notes is a very poor idea because you may later wish to know what you had erased. A single line through incorrect notes conveys the same meaning and leaves all notes legible.



Field Notes


At the start of each observation session, you should note the subjects' physical environment (e.g., wind velocity and direction, temperature, percent cloud cover).  Sketch a map of the study area emphasizing components influential to your work.  Throughout each session, keep track of the time of day.  Glance at your watch periodically and note the time in the left-hand margin of your notebook.

Behavioral data should be recorded truthfully and without preconceptions of what an animal should be doing.  In modern ethological studies, behavior must be quantified.  Phrases like “several times a minute” or “a few meters”, have no place in field notes.  Using the metric system, always put number estimates on behavior; the whitetail deer walked 7 m in 5 sec, or the pigeon raised its wing 12 cm.

Field notes (and your thinking while recording them) should be couched in operational terms.  Simply record what you see without ascribing motive or function to the behavior.  This is an important and sometimes quite a subtle skill, but one you should cultivate.

Incorporate in your notes frequent drawings of behavioral patterns; stick figures are perfectly adequate.  Besides giving you a pictorial record, drawing forces your attention to all aspects of behavior.  Only in drawing the posture of a drinking gray squirrel (Sciurus carolinensis) might you realize that you do not know the position of the tail.  A camera can be a valuable (though not required) tool to aid analysis and, later, illustration of the ethogram.



The Ethogram


An ethogram is a distillation of field notes.  It involves the grouping of similar behavioral patterns shown at different times by one or more species members.  That is, categories of behavior are identified and described.  To some degree, functional interpretation must come into play here.  For instance, collecting a constellation of behavioral patterns under the rubric, hunting behavior, requires the assumption that all the behavior so labeled has food-finding as its purpose or function.  Some items in an ethogram are relatively free from such problems.  The flight of birds can certainly be safely considered under the category, locomotion.

A good plan for organizing your field notes into an ethogram is to look for a hierarchy of categories and subcategories.  Someone studying mallard ducks (Anas platvrhynchos) might consider locomotion as one major category.  This could then be divided into flight, swimming, and walking.  Flight could be split into flapping and non-flapping, swimming into surface and subsurface.  Flapping might be further dichotomized into moving and stationary (hovering), and so on.  Although I encourage you to structure your ethogram however you wish, other major categories you might consider (always being cautious in ascribing function to behavioral patterns) are drinking, feeding, hunting, bodily maintenance (grooming, sharpening claws, etc.), sleeping, resting, courtship, and care of young.

Your ethogram should be quantitative, and should attempt to describe variation in behavior.  Field notes can be converted into averages, ranges, and even standard deviations or coefficients of variation.  On average, how many times in succession do mallard ducks preen their breast feathers?  Do males and females differ in this?  Is there more variation among individuals of one sex than the other?

When examining and synthesizing field notes, be aware of what you did not observe, what the species did not do.  For example, there is a medial region of the dorsum that house sparrows cannot reach with bill or feet.  This apparently is the only area the sparrows never preen.

Ethograms benefit greatly from plentiful illustrations (stick figures, line drawings, sketches, and/or photographs) complementing the text.  Even ethologists with very little artistic talent have no trouble tracing line drawings from photographs, the most prevalent tactic in published ethograms.



The Formal Write-up


Scientists must be able to write well, not necessarily with flair, but accurately, efficiently, and often within rather strict guidelines (at least when writing for publication in technical journals).  I wish you to experience writing an organized report in a form suitable for publication.  A scientific paper is divided into several sections; all must be typed with double-line spacing throughout.  Below are more detailed descriptions of these components of a publication.


Title page: As the name suggests, this is a single page delineating the article's title and the author's name and address.  If a species' common name appears in the title, the scientific name should follow immediately after.  However, the scientific name may be used alone.


Abstract: Any given journal will require either a terminal summary or an initial abstract, of about 5%, or less, the length of the entire report. I ask for an abstract that will follow immediately after the title page.  The title of the paper should be repeated at the head of the abstract.  Present scientific names of species after first use of their common names.


Introduction: Beginning on the page after the abstract, this section generally presents theoretical background followed by hypotheses and predictions leading to the present work.  In your ethogram, this section may be short, perhaps limited to describing the taxonomic status, geographical distribution, trophic level, etc., of the study species.  The title should also head the first page of the introduction.  Present scientific names of species after first use of their common names here and throughout the rest of the ethogram.


Methods: This important section should describe how, when, and where you worked in language precise enough to allow replication of your study by another investigator (the essence of the term, re­search).  Present information on study site, temporal intervals, recording techniques, and equipment.  Analytical procedures, if any, also belong here.


Results: The heart of the paper, this section lays out in organized fashion what you have discovered.  How you interpret what you have found should be separated from Results, and left to the following (Discussion) section.  In your results section, I expect to find the annotated inventory of behavioral patterns condensed from your field notes, along with references to illustrative tables and figures included in your report.


            A few words about tables and figures.  Each table should appear on a separate page, be numbered with an Arabic number (not the page number) so it can be referred to easily in the text, and .be headed by an accompanying legend (actually the title of the table).  In a scientific report, all graphs, drawings, photographs, diagrams, etc., are called figures. 

            Each figure should also occupy a separate page, have a legend (at the bottom of the page), and be given an unique Arabic number.  Refer to tables and figures in the text in the following fashion: “Horses used two types of trotting gaits (Fig. 8), one was contralateral ...” or “... while male cardinals(Cardinalis cardinalis) sang for longer periods than females (Table 4)...”


Discussion: This section immediately follows Results.  Here, should appear your interpretation of the behavior you recorded.  You may wish to discuss possible reasons for variation of behavior with the same apparent function between sexes, among different age groups, or under differing environmental conditions.

            You may have noticed that I have said nothing about your referring to the literature in conjunction with this project.  I wish you to delay any literature search until you have finished your field work and written the Results section of your ethogram.  Thenceforth, you may scour the literature for works related to your project.  Here, in the discussion section, is the place to compare your results and interpretations with previously published findings.  Works referred to should be cited parenthetically in the text by author or authors (last names only) and date of publication.  For example, “Wilson (1968) reported that...”, or “ ... mourning doves (Zenaidura macrura) court by cooing (Brown and Jones 1972).” Publications with more than two authors should be referred to in the text by first author's last name followed by et al., e.g., (Smith et al. 1973).  Notice that et al. is an abbreviation for et alii (Latin for ‘and others’), so that al is followed by a period.  Several papers cited together should be listed chronologically by date of publication (not alphabetically by author's last name) and separated by commas, e.g., (Lack 1944, Morse 1968, Balda 1972).


Acknowledgments: Immediately following the discussion section, this section details those who have helped you, people who have loaned equipment, granted permission to use a study area, criticized . your manuscript, and so on.  In major studies, long-suffering significant others often are publicly thanked here.  Those fortunate to have financial support for the study acknowledge that here, also.


Literature Cited: This, the final section of the report, lists in alphabetical order of first authors, last names only those works actually cited in the preceding text.  In typing this section, place the first line of each reference at the left-hand margin and indent succeeding lines.  For journal articles, the usual procedure is to list in succession the author's surname, then other initials, the date of publication, title, journal name, volume, and page numbers.  Contrary to the discussion section, here all authors of multiple-author papers are listed.  For books, the title is followed by the edition number if not the first edition, then the city of publication and the publisher's name.  Sometimes a paper will appear in a book edited by a person other than the paper's author.  This should be referenced in the manner of the next to last entry in the Literature Cited sample presented below.


Campbell, A. 1965.  Whooping cough in cranes.  New York, Academic Pirates Press.


Mallock, W. B. J., Wilson, C., and Goldstein, A. 1975.  The courtship behavior of Homo sapiens.  J. Comp.  Hanky Panky 17:532-541.


Tell, W., Jr. 1862.  A close call. in The aerodynamic characteristics of crossbow bolts.  W. Tell, Sr. (ed.), Zurich, Apple Cider Press.

Warlock, W. 1862.  Which witch is which?  J. Salem Follies 1:23-25.

Miscellaneous addenda


Pagination.  The title page is unnumbered.  The abstract page or pages should be numbered consecutively beginning with page 1. Begin a new page with the introduction and give it the number following the number of the last abstract page.  No succeeding section of the text should start with a new page; sections should be separated only by section headings.  Tables should appear in internal sequential order behind Literature Cited, each table given a separate page.  Finally come the figures, each given a separate page.  To summarize:


Title page:     unnumbered.

Abstract:         new page; numbered consecutively beginning with page 1; head first page of abstract with paper title.

Introduction, Methods, Results, Discussion, Acknowledgments, Literature Cited: begin introduction with the title; pages numbered consecutively; sections separated only by section headings.

Tables:           each given a new, consecutively-numbered page.

Figures:          each given a new, consecutively-numbered page.


Scientific names.  Immediately after first mention of the . common (usually English) name of a species in the text, the species’ scientific name should follow in parentheses, as we have done in this paper.  Thereafter, the common name may be used alone.  Scientific names are always underlined or italicized in typescripts.


Units.  Metric system units should always be abbreviated in the standard fashion (e.g., m for meter, cm for centimeter, sec for second, g for gram, etc.). I ask you to avoid English units if possible, but if you must use them, follow the English unit with the metric equivalent in parentheses, '...sat within in inch (2.5 cm) of its mate.'


Numbers.  When a number is followed by a unit of measurement, the numeral is always used (5 days, 8 feet, 3 sec, 7 m).  When a number is not followed by a unit of measurement, it is spelled out if it is one digit, or if it begins a sentence.  If it is at least two digits and does not begin a sentence, the number is given in its digital form.  Thus, “Twelve whitetail deer were...” “The study area measured 22 by 36 cm...”, and “Downwind of the lake, four mallard ducks were...”


Dates and times.  Dates are written in the form 17 April 19xx.  Use the 24-hour clock to report times (e.g., 09:52, 13:15, 22:57).


Grammar and Spelling.  While I have no formal guidelines for penalizing poorly-written papers, I expect your paper to be well-written.  If your grammar and/or spelling are not good, I expect you to use a spell-checker if your software has one, and then to have someone edit the preliminary draft of your paper.