Professor in charge:
BSCI 465 / Biol 708D
COURSE INFORMATION
Fall 2013 M-W 10:00-11:15; 1117 PLS
Dr. Gerald Borgia, Biology Department
Office hours: After class Wednesday and by appointment
Office BP 4239 Email Borgia@umd.edu
Living organisms had existed on Earth, without ever knowing why, for more than three
billion years before the truth finally dawned on one of them. His name was Charles
Darwin. – Richard Dawkins, 1976
The question "What is man?" is probably the most profound …[and] … all attempts to
answer that question before 1859 are worthless and that we will be better off if we
ignore them completely. —Simpson 1966, p. 472
Overview
Welcome to BSCI 465/Biol 708D, Behavioral Ecology. In this course you will
become acquainted with the some fundamental principles of evolutionary biology and
how they can be applied to study current issues in animal and human behavior. This
course is directed at answering the question: Why do living things do what they do?
We will be interested in how behaviors evolve and in determining the functional
significance of behaviors in all organisms. Issues associated with behavior present
some of the most significant challenges to evolutionary theory because of the
complications associated with interactions between (and within) individuals that have
different reproductive interests.
For many years the study of animal behavior had no theoretical focus and many
important and interesting questions went unanswered (or incorrectly answered)
because biologists working in this area did not have an adequate theoretical model to
deal with them. Much of this changed with the integration of modern evolutionary
theory with the animal behavior and has let to the advent of behavioral ecology and
sociobiology (to most biologists these are interchangeable labels). We will, as a
consequence of this evolutionary focus, deal with a much broader definition of behavior
than is commonly used in animal behavior courses. We will be as interested in the
behavior of chromosomes as we are in the parental care of honey bees, or in human
mate choice. We will focus on why certain behaviors and related traits evolve with
particular emphasis on difficult evolutionary problems. Some questions we will consider
are: Does altruism evolve? How do conflicts of interest arise and shape social
interactions in mate choice, social grouping and meiosis? Why do certain adaptations,
such as sexuality, that greatly affect social relationships in nearly all species, exist?
One of the most useful techniques for studying evolution generally and behavior in
particular is to study adaptations. Adaptations are traits that show design for
reproduction of the organism carrying the trait. Natural selection is the key force
shaping new adaptations and the ongoing pressure that it exerts maintains organisms
in a reasonably well-adapted condition. Natural selection favors the reproduction of
variants that are best able to deal with immediate environmental and social
contingencies. The cumulative effect of this selection is to favor the reproduction of
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genes that produce these effective “tools” against genetic alternatives that are less
effective. To the extent that these tools (or traits) are inherited and, within the context
of evolutionary time, organisms track their environment, producing and “perfecting”
adaptations that have proven useful to their reproduction.
The dominance of natural selection causes us to predict a close fit between the
environment and the kinds of adaptations carried by an organism. This prediction is
very useful for several reasons. First, it allows us to predict what kinds of behaviors
(and, more generally, adaptations) we expect to see (and not see). Second, where
there are deviations from these predictions there is often evidence of interesting and
previously unnoticed theoretical issues. Third, a high level of adaptation causes us to
look for functional significance where there is evidence of complex design. Fourth,
complex adaptations are built up over time and as such their presence provides
evidence of the past history of natural selection. We integrate these ideas with modern
population genetics and to produce a model of behavioral evolution. We can then
explore patterns of adaptation to test this general model and understand more specific
and difficult cases of behavioral adaptation.
Course objectives:
1) To familiarize students with important issues in behavioral ecology, broadly defined.
2) To teach students “evolutionary thinking” which provides a powerful tool for solving
problems in all of biology.
3) To critically consider scientific hypotheses and related tests in the literature.
Course organization:
BSCI 465/708D are 3 credit courses divided into two one hour and fifteen minute
sessions per week. Monday will be a full 75 minute lecture and Wednesday will be split
between lecture and discussion of assigned papers. Students are required to read
papers in preparation for the discussion.
Your final grade will be based on a total of 500 points. There will be one mid-term
exam (100 points), a final (150 points/cumulative), one essay (135 points – 100 for the
exam and 35 for meeting deadlines in paper preparation) and discussion (115 points).
Since grading is on total points, a poor showing in one part of the course could result in
a poor grade for the entire course. To guard against this possibility, it is important that
you be an active participant in all phases of the course. Students are expected to
follow University Honor Code and follow University policy for absences.
Essays are expected to provide critical discussion of an idea or how an idea has
been treated by another author. Essays should be 2 to 5 pages in length, and must be
typewritten and double spaced with a few, usually not more than 5, relevant references
You should write in the style of a scientific journal, with a concise style and include the
points that you believe that are needed to explain the idea you are developing.
References should follow the format of the journals Animal Behaviour, Behavioral
Ecology or Evolution. Essays should be submitted electronically as an attachment in
MS word doc file to Borgia@umd.edu with your name in the file name and you email
address in the paper header.
I require that the topic and outline of the essays must be discussed with me
and approved before the essay is submitted (no emails to discuss please). This
commonly entails a student finding a topic he or she wishes to write about and then a
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brief visit with me to discuss the topic (By November 1; 15 points). There should then
be a second visit in which we a written outline developed by the student that indicates
the flow and logical sequence of the arguments to be presented in the essay (By
November 9; 20 pts.). Paper or book summaries are not suitable, although critical
reviews that offer a significantly different perspective and or analysis of an idea or
viewpoint are. The key point is that I am looking for am original intellectual contribution
from you. Suitable paper topics should involve issues discussed in the course or which
are related to the theme of the course. The goal here is not to recycle old term papers
or thesis proposals. I will provide lists of paper topics well in advance of the deadline.
Essays are due November 15; 100 pts) unless we agree on an alternative date.
Because essays require discussions with me please do not wait until the last minute
to make appointments to discuss papers.
Readings:
Readings for this course are based on the primary literature. I will post materials s
the week before when the reading is due. Students are expected to have read the
material and be ready to actively participate in class discussions based on the readings.
A significant part of the course grade will depend on student preparation and
participation in course discussions (115 points).
For extra help the book The Selfish Gene (second edition) by Richard Dawkins is
highly recommended as a helpful overview and guide to how issues are thought of in
this course. Other useful books include Principals of Social Behavior, By Andrew
Bourke, Behavioral ecology by J.R. Krebs and N. Davies, Adaptation and Natural
Selection by George Williams, and Sex, Evolution and Behavior by M. Wilson and M.
Daly. References to additional readings will be provided as the course progresses.
Special notices:
1. Evolution is treated as a fact by all leading biologists and evolutionary theory is the
central theoretical construct of biology. This course is centered on the view that
evolution by natural selection has been critical in shaping life on earth including human
behavior. While students are not required to believe that evolution has happen they
should be warned that to be successful in this course they will have to use evolutionary
ideas in all phases of the course.
2. Students should note that the evolution of sex and sexual tactics in animals and
humans is a major component of this course.
3. To be successful in this course it’s important that you regularly attend lecture,
take detailed notes and actively participate in discussions.
Special considerations
Academic dishonesty is incompatible with good science. As presumptive scientists I
hold you to the standards of our discipline ergo cheating is not tolerated. Note that the
University of Maryland sometimes treats academic dishonesty as a serious offense so it
is in your interest to be familiar with the relevant sections of the student handbook.
Make-up exams will not be given except in cases for which there is an appropriate
medical reason.
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BSCI 465/Biol 708D Lecture Schedule
Tentative Lecture Topic
Introduction: Darwin, Evolution and Behavioral Ecology
Natural selection and other agents of selection
Adaptation : Levels of selection, selfish genetic elements, and
inclusive fitness
Levels of selection: Why lemmings don’t jump off cliffs
Evolution as a predictive model: Adaptationism
The debate over adaptationism: The clitoris debate
Selfish genetic elements
Gamete dimorphism and Bateman’s argument
Evolution of mating systems
Sexual selection models I: Runaway, Good genes, preexisting preference
Sexual selection models II: Cooption and preexisting traits (bird song)
Sperm competition, genetalia and male-female conflict
Why do bowerbirds build bowers? Sexual conflict and cooperation
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14.
Midterm Exam
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15. Sex ratio: Inbreeding and outbreeding models
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16. Evolution of sex
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17. Inclusive fitness as an ESS
Nov
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18. Selection in small groups, group selection and altruism
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19. Helping non-relatives -- reciprocity and cooperation
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20. Memes, NS and the role of learning
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21. Social evolution in insects and reproductive division of labor
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22. Sociality in birds
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23. Mammal social organization
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24. Social structure in primates and hominids
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Sexual selection in humans
Dec
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26. Human sexual conflict: MGM and FGM
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Human brain and social evolution
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28. Cultural evolution / Darwinian and Evolutionary Psychology
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29. Darwinian medicine
Final: Saturday, December 21, 8:00-10:00AM, Rm 1117 PLS (No early exams)
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Discussing and critiquing scientific papers
A major portion of this course deals with the discussion and criticism of scientific papers. In
an ideal world all scientific papers would be clearly and concisely written, consider and
effectively test all reasonable alternative hypotheses, have a large data base on which to draw
conclusions, use appropriate statistics, be free of errors and, most importantly, state
conclusions that are supported by the results. While many papers do meet these criteria many
others do not sp scientific papers must be read (and written) from a critical viewpoint if they are
to accomplish their goal of advancing our knowledge of biology. Almost all scientific papers
undergo some review before they are published but the level of scrutiny is highly variable so
even though they have undergone review papers with major errors are often published.
No study is perfect but we can establish reasonable standards of what to expect. Standards
change and almost always in the direction of increased stringency as a field develops. These
may occur for a variety of reasons including a willingness to accept more questionable
information when a field is new, the development of new and more accurate (and usually more
complicated) techniques for analysis as a field develops, and increased competition for journal
space. Regrettably many studies are considered important because of they were early in the
development of a field, or because of who did them and not how well they are done.
It is difficult to do good science and present scientific results papers well so we need not
assume that papers with weaknesses are done with bad intent. It may be physically and
economically impossible to collect all the desired data, and there are limits to the amount of
time that can be devoted to doing analyses and writing the paper. What is amazing is the
number of quality papers that are written despite these and other limitations. We should,
however, require that authors meet reasonable standards and are straightforward about the
limitations of their work. Part of this evaluation process comes from your experience in reading
papers so that you can appreciate the range of variation in the quality of papers. Below are
some questions you should consider in evaluating the papers you read.
What questions does the publication address?
Is there a significant question being addressed? There is a nearly endless number of
questions that can be studied so it is important to ask if the questions addressed are of general
interest. How do we know if a question is important/general?
Does the study tests hypotheses that address important theoretical issues? We develop
major hypotheses or theories to provide and understanding of how the world works. Generally,
the more the hypothesis explains the greater its importance, e.g. does it explain or improve our
understanding of some critical process in animals and/or humans? In this course we will value
papers based on their ability to clarify issues important to understanding evolutionary and
behavioral models. Papers that simply describe a particular kind of animal behavior are
considered less valuable than those that use information about behaviors to develop and test
general hypotheses about why that behavior occurs, particularly if that behavior is unexpected
based on existing theory or if a there is new theory that has not previously been supported.
Does the issue raised address a real problem or is it a result of unclear thinking?
Statements may seem important because they have been linked to important processes but
closer inspection shows this linkage is in error. Is the question falsifiable? Will any result
support a particular conclusion or are the alternatives spelled out clearly.
Does the publication address the question in a reasonable way?
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Does the study make reasonable assumptions? No study can start from scratch and work
must be done in the context of is already known. Does what it assumes as a starting point
generally fit within this knowledge? This can be a problem for truly revolutionary studies
because they often don't fit. The assumption of a paradigm for a field has the advantage of
helping you sort through lots of information but if it is not reliable important discoveries that do
not fit the conventional wisdom may be ignored. Every scientist must deal with what they
believe to be realistic and not and this is often only obtained with experience.
Are all reasonable hypotheses considered and tested in some way? There are often many
possible explanations for a set of observations. We must make assumptions about what
hypotheses we reasonably need to test. We must ask if an author has been fair in choosing
what hypotheses to consider. Often not all hypotheses can be considered. Has the author
been forthright in explaining what has not been done.
Is the general question that is presented actually addressed by the experiments and or
observations?
Our ability to judge the significance of a study depends on the author’s ability to properly set
the context for existing theoretical and empirical information. Does the author give a clear and
fair (not inflated) picture of the meaning of the study and what has been done by others?
For theoretical papers does the model accurately reflect natural systems? In biology, as
opposed to mathematics, models are not of interest because of their intrinsic properties but
because they provide accurate representations of the real world. Often reality is complex and it
is necessary to simplify. Do the assumptions the model makes allow it to still provide a fair
representation (one that can be tested empirically) of the real world. Are the simplifications and
the domain over which the model is robust clearly stated?
Data collection and treatment
Most studies are done to make inferences about a larger set of organisms than were
actually studied and to help resolve problems beyond those addressed in a particular
experiment. Are means used to make these inferences reasonable? For example, are the
target organisms of the study appropriate to the issues being considered?
Are the study populations reasonable? Are they representative of the population they are
supposed to be making inferences about or are they in marginal or disturbed habitats? This is
particularly important for evolutionary studies because it is often assumed that current patterns
are representative of past conditions (when the traits under study evolved).
Are experiments reasonable? Do they ask the animal to do something it can't do? Are they
properly controlled? Do they ignore factors like experience that isn't or can't be controlled.
Do the variables measured actually measure what they claim (or what their name
suggests)?
Are there results not presented that can alter conclusions? It is important to present
negative as well as positive results.
Is there data that should have been collected that could clarify or alter conclusions?
Do the conclusions follow from the data? Are all reasonable interpretations considered? Is
there too much effort directed at explaining away inconvenient facts?
Are there hidden biases in the data collection. Is observer bias considered and dealt with?
Are the results of multiple tests "factored in" the assessment of statistical significance?
Scientists can go “fishing” and if they do enough tests some will be statistically significant by
chance.
Do the statistics procedures support the rules of statistical inference? (Unless you have had
a statistics course or two you probably wouldn't know too much about this!). Especially
important are the issues of independence, unintended/hidden correlations, small sample size,
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and use of appropriate tests (including meeting assumptions of tests).
Are graphs and other figures understandable and properly labeled? Are all symbols
defined?
Do the summary and/or conclusions accurately reflect what was found in the body of the
paper?
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