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Field Studies in the Evolution of Animal Behavior

Summer 2024 at Flathead Lake Biological Station

Northwestern Montana

This is an intensive 4-week, 5-credit course offered during the second half of the 2023 Summer Session at Flathead Lake Biological Station in the magnificent Northwestern Montana Rockies; July 1- August 19. Students who desire extended research experience and exposure to conceptual material may opt to arrive 2-4 weeks early, signing up for a seamlessly integrated 2 or 4 weeks of Advanced Undergraduate Research supervised by Dr. Watson; June 17 - July 12. Committing to the additional 4-week will give you a better taste of the whole scientific process. But, however long you attend, this program of study is designed to prepare you for graduate work in Evolutionary Behavioral Ecology and Psychology. It is important to note, however, by its nature and design, my animal behavior courses also are of great interest to Social Science, Liberal Arts, or Humanities majors, including budding Psychologists and Philosophers.

Here is the FLBS 2024 Summer Session Web Site:

For more information about enrollment reach out to Hannah Gerhard, the Biostation's summer session manager.

Please also feel fee to contact me at with questions about course content!

Do not neglect Biostation scholarship opportunities!

The deadline for scholarship applications is early March.

Please do not sign up and hold a spot for yourself in this course if you are not committed or financially prepared. Slots are limited, and holding one may prevent others from signing up. Late withdrawals can cause the course to be canceled; this has happened and it's a bummer.

Transferable undergraduate or graduate credits are awarded by the University of Montana (UM), Missoula, MT, USA.

Students of evolution and behavior from across disciplines and around the world are invited to participate; maximum enrollment will be 6-12 students. Most of my students are biology majors, but as in years past I enthusiastically consider applications from upper division and graduate students in the humanities who have a deep desire to understand the evolution of behavior. I once had a philosophy Master's student from American University, Beruit, Lebanon, who with the help of my FLBS behavior course went on to do her doctoral work in Evolutionary Anthropology at Rutgers.

Aims of the Course
Our big question in this course is always how are animal minds designed to solve niche-specific ecological and, ultimately, reproductive problems? That is, problems that affect each individual's nonconscious prime mandate of maximizing lifetime inclusive fitness.

Behavioral ecology is the study of animal minds, their functionally related morphological and physiological traits, and the relation of their embodied mind's design features to historical, recent and currently observable forces of natural selection. Evolutionary behavioral ecologists study how natural selection in all its forms has fashioned wild minds to gather, process, transmit, and respond to streams of information coming from within the body, via self-assessment, as well as from the environment. In this class we practice and discuss how to make evolutionary sense of the spectacular and often puzzling design features of animal behavior, and their awesome responsiveness to complex variation in their natural environments. Moreover, I emphasize going for deep connection, real contact, with the animals you observe, and with yourself as you observe them.
 I live at the station and am available to all my students 24/7 during the summer session!

Flathead Lake Biological Station has been the main site for my research on the sierra dome spider since 1980, the year I first took this very class myself. I now teach this course in an updated form that still honors the approach of my wonderful professor from that auspicious year, the late Dr. Allen Stokes, along with the cutting edge approach to field work of my superb doctoral and postdoctoral mentors Stephen Emlen, Paul W. Sherman, William D. Hamilton, and Randy Thornhill at Cornell, Oxford, and the University of New Mexico, respectively. I assisted in teaching rigorous field courses in animal behavior at Cornell University for two terms under Dr. Paul W. Sherman. I have formally and informally mentored many undergradute and graduate students. I have taught Field Studies in the Evolution of Animal Behavior at Flathead Lake, in both 4-week and primarily 8-week formats, since 1994. Thanks to the revivification of the FLBS summer teaching program after a hiatus and Covid-19 delays, we successfully reinstituted the class in a new 4 to 8 week format in 2023.

The course is designed to provide mature, persevering organismal biology undergraduates and graduates, teachers, and select students from other fields of behavior (e.g., psychology, anthropology, philosophy and the humanities) with substantive experience in performing professional quality research in evolutionary behavioral ecology. Necessarily then, the course also is intended to hone one's ability to think creatively and productively from an evolutionary "adaptationist" perspective. In my experience, it is only through concentrated contact with diverse animals operating in nature, combined with on-the-spot discussions of the often profound evolutionary principles germane to understanding their behavior and related morphological and physiological design, that most people really begin to appreciate the grandeur of natural selection and the staggeringly beautiful creatures it has produced. Likewise, it is only via thorough, determined digestion of evolutionary concepts and direct involvement in designing and implementing rigorous research methodologies, that one develops and understanding of the power of evolutionary biological thinking for elucidating the structure and functional significance of animals' bodies and minds.

Students who immerse themselves in the course will be prepared to undertake creative doctoral quality research in behavioral ecology. Students often also have had their world views transformed and enriched, sometimes radically. This is not so much my doing. Rather, I think it is the nearly inevitable combined effect that (1) sustained, disciplined contact with nature and (2) sustained, disciplined exposure to correct theory, has upon perceptive truth-seeking human minds. Sadly, few people ever have an opportunity to work with others, to seek a richer more objective understanding of mind and nature, under these two key synergistic conditions.

The course is intended for students who wish to move beyond the usual undergraduate role of knowledge consumer and into the role of the active scientist: a collaborative creator of new verifiable knowledge. The course's top priority will be the development of practical and analytical skills gained while performing collaborative, theoretically informed and potentially publishable field and lab projects. Evolutionary analysis of behavior will be emphasized together with techniques for capturing, handling, marking and observing animals in naturalistic and experimental contexts enabling testing of hypotheses about the behavior's adaptive function. Student projects will include all elements of the research process from initial observation of a behavior, to brainstorming about its possible adaptive functions and methods to study them, to statistical data analysis and writing up the project's results in publishable format.

Studying adaptation is surprisingly tricky, but tremendously intriguing and rewarding. The appropriateness and efficacy of the three basic modes of investigation - the comparative method, measurement of fitness differentials among behavioral variants, analysis of functional design - are debated in the literature. These methods and their associated problems and controversies will be introduced with the aid of recent papers from the primary literature. The student's digestion of such conceptual material will be facilitated by practical experience in: (1) mentally carving complex animal phenotypes into adaptively meaningful suites of behavioral and morphological traits, (2) developing evolutionary hypotheses about the costs and benefits and potential adaptive function(s) of observed behavior, (3) choosing among available modes of investigation to establish formal testing schemes designed to critically evaluate appropriate alternative hypotheses. By the way, many of the most intriguing and observable creatures to study are arthropods. You might want to think twice about taking this course if you have a strong bias against studying watching and possibly handling them. On the other hand, you may discover a wonderful new appreciation of them in this class. I certainly never thought, back in 1980, that I would spend over three decades studying the sexual selection system of a spider, but it is one of the best things that ever happened to me. Attentive observation coupled with evolutionary thinking will be the foundation for everything we do in the course. Students will learn how to combine observations of an animal's behavior and ecology with knowledge of natural selection theory to develop alternative evolutionary hypotheses and testable predictions as the basis for their research projects. Students will be thoroughly coached in the computerized statistical analysis of their research data. Writing skills will be honed in 4-5 essays on theoretical topics and in the production of at least two research papers in which the quality of writing and the integrity of the student's collaborative effort will play a substantial role in grading. Patience, fortitude, concentration, adaptability and the ability to distinguish simple perception from subtly biased interpretation will be some of the "inner skills" that students will be invited to work on during the course.

In all aspects of the course, the student's initiative, originality will be encouraged and their observing and questioning faculties will be honed. Yet, detailed guidance also will be available. Lectures and discussions as well as extensive reading will provide students with the essential theoretical knowledge to develop and interpret their research projects from a modern evolutionary perspective. According to the classes' interest, we shall also bring evolutionary psychological perspectives on human behavior and the design of human minds into the picture via reading, critical discussion and, perhaps, self-observation.

Primary responsibility for each research project will rest with teams of 2-3 students: the "Principal Investigators" (PI's). Each student will serve as a PI on two projects. PI's will be co-responsible for (1) developing an especially thorough understanding of the conceptual and logistical aspects of their project, (2) analyzing and writing-up the results in publication format and (3) giving a 20 minute end-of-semester oral presentation in collaboration with his or her co-PI's. However, every student in the course will be involved in the development and troubleshooting phases of every project via research roundtable discussions. All students also will assist in data collection in all projects on a rotating basis. Scheduling of each student's participation in data collection will be handled by project PI's in coordination with the instructor. A major component of the course grade (50%) will hinge on the student's engaged participation in the research projects in both their PI and research assistant capacities, as well as their thoughtful contribution to class discussions. A number of class projects involving highly observable and manipulable arthropods and other critters have been planned in some detail ahead of time, in the hope that all students will have the experience of participating in successful research with publication potential even in the short time available for the course. However, students also will have opportunities to discover and develop projects of their own.

Prerequisites and Recommended Prior Courses
Instructor approval is the only prerequsite, because I wish to teach evolutionary theory and a modern Darwinian approach to understanding mind and nature to any serious upper division undergraduate or early career graduate student. Science teachers are also welcome. All students should directly consult the instructor before enrolling. Some of my best students have been Humanities majors. I work with every student according to their background and am dedicated to providing you the background you need to thrive in the course.

Course Format
As a course that centers on real research on real animals, the format must be dynamic and flexible. Nature does not care about anyone's plans or wishes to order their work in some comfortable fashion. We must be flexible and opportunistic or we will miss many of our best chances for research. The class needs to be a rapid-deployment-force!

Willingness to participate in a professional and collaborative manner is required during official class hours. Additional work is encouraged during the evenings and at other times of the week as the student's schedule permits. Students in charge of caring for captive animals will be expected to care for them meticulously throughout the week and to arrange for another student to take over this responsibility on days when this is impossible.

I will be living at the station all summer, enjoying meals a the commissary, and will be available to you for research support and all manner of course-related discussion during the entire week. I love my work and this course and am hopelessly afflicted with an overwhelming desire to assist and guide curious motivated students!

More on the Design of the Course
Field Trips - Although there will be some field trips, you should know from the outset that this course is not designed to be a tour of NW Montana power-spots, as wonderful as this would be for all of us. Rather, it is a course in which attempts to do meaningful behavioral research is our first priority. For the sake of practicality, most of our time will be spent at or near the station.

Research - The research experiences provided in this class are not "exercises." They all require genuine innovation both in the formulation of hypotheses the development of research methods. Moreover, the kinds of questions our class projects address are of current interest to behavioral ecologists and the write-ups should have high publication potential if we, as a team, obtain sufficient high quality data. With good fortune and hard work this should be feasible in many cases, although it will never be easy. Examples of class projects include:

  • Social behaviors of Columbian ground squirrels. In 2019 I successfully reintroduced these intriguing creatures to the Biostation grounds. (The original native population was wiped out by a family of mink in 2004 and never recovered.) They have spread across the station and are thriving. Many are microchipped and of known relatedness setting us up to examine substantive hypotheses concerning their patterns of social behavior. All students will be assigned 1 or 2 ground squirrels to get to know as closely as possible during the course.

  • The effect of operational sex ratio on the intensity and duration of intersexual conflicts over mating duration in two water strider species (Gerris buenoi & G. incurvatus.)

  • Comparative female mating resistance in water striders in relation to genitalic morphology.

  • Violent male-male competition and chemical communication in a lekking micro-lepidopteran moth (Adela sp.)

  • Female nutritional status, mate number and mate choice in mormon crickets. (Anabrus simplex)

  • Correlates of territory size and shape in Sorex shrews.

  • Fluctuating asymmetry, lek attendance and sexual success in mayflies. (Hexigenia sp.)

  • Colony life of Lasius sp. ants, employing large plaster glass-sided ant farms with outdoor access for the ants.

  • Fighting ability and copulatory performance (i.e., sexual competitiveness) of male sierra dome spiders (Neriene [=Linyphia] litigiosa) in relation to rates of aging.

  • Sexual division of labor in nesting birds.

  • The adaptive function of male mimicry by female damselflies (Enallagma boreale).

  • The function of low nectar yield aphid guarding in several ant species (e.g., Campanotus pennsylvanicus) - possibly an unusual example of chemical exploitation of ants by aphids, or ants investing in small colonies of aphids so that they can grow and eventually provide profitable amounts of honeydew for the ant colony (i.e., farming).

  • Sex pheromones, female vibrational signals, and fighting intensity in male sierra dome spiders.

  • Nutritional status and sexual receptivity in a water strider (Aquarius remigis).

  • Comparative foraging behavior of various ant species.

  • The intensity of male-male courtship and competition in sierra dome spiders in relation to female quality, relative and absolute male sizes, breeding season date, air quality, etc.

  • Interspecific and intersexual competition and nectar nutritional content in Rufus and Calopie Hummingbirds.

Alternative student-initiated projects may be performed by individuals who show strong ability and desire to work independently or by teams of 2-3 students (the latter is encouraged). Student-initiated projects may be done in place of one or both of the projects listed above. During the first two weeks of class, students will be encouraged to search the station for potential research projects. There are many species with interesting behaviors awaiting discovery and study! Student- initiated projects may consist of any research scheme that a student can devise. After preliminary observation of the animals and behaviors in question, the student who wishes to attempt an original project should meet with the professor and intern to discuss the feasibility and design of the research. After receiving the instructors' input, the student must settle on a behavioral research question and a research protocol and have the final project approved by the instructor. If the research is approved, the entire class will meet to discuss the ideas involved, after which the relevant student(s) will be turned loose on the project. The main criteria for deciding whether a proposed project can be approved are that a specific evolutionary hypothesis must be tested concerning a behavioral or morphological trait of a readily observable animal that lives on or near the biological station. Students performing projects of their own design may be excused from certain whole-class activities in order that they be able to devote time to their projects. A student with a good independent project may be excused from prior commitment to lead a pre-planned project within the first 4 weeks of the course. We will do everything in our power to facilitate performance of approved student-initiated projects.

Lectures and discussion 

I believe in teaching through conversation, although some straight lecture will be necessary early in the course to efficiently introduce central concepts. We will try to concentrate lecture/discussions into times of poor weather or other times when observation of animals is likely to be less productive. We shall concentrate most lectures into the first several weeks of the course, so that there will be more research time in late June, July and August when weather is more likely to be permissive. Lectures will be scheduled on a varying basis to maximize opportunities for research activities. Usually, there will not be more than 1-2 hours of lecture on any given day, except perhaps the first few class meetings. Again, the main point of the course is to get practical field experience!


Students will maintain notebooks not just of class notes, but ideas and musings related to course ideas, which may be brought up in class during daily discussions. I will propose thought questions for you to ponder, individually and together,  and write succinct passages on. Writing is an important thinking tool! I will evaluate the thought and effort put into your notebooks as part of your grade.

Students are encouraged to confer with one another and with the instructor in analyzing the assigned topic, but individuals must write semi-final drafts of their essays completely independently. Come up with your own way of wording whatever you decide to present. As you confer with your peers or instructors, try to be subject to logical argument without letting your originality get squelched unless there is a compelling reason! There may be more than one approach to dealing with any assigned question.


Oral Presentations 

So that the entire class may enjoy the fruits of their data collection and project planning efforts, team leaders of each fully-planned and exploratory project, as well as any independent project, will present the results of the project to the class. Presentation of the results of exploratory projects (about 15 - 30 minutes long) will be in the evening within 2 weeks of the execution of the project. Presentation of fully-planned project results (20 minutes long) will be anytime during the final two weeks of class.

Reading and Discussions 

Students will read mostly self-assigned portions of the deeply hallowed course textbook, John Alcock's, "Animal Behavior: An Evolutionary Approach" in its beautiful 10th, 11th, or 12th edition, now co-authored or sole-authored with/by Dustin R. Rubenstein. Specific readings from the text will be chosen by each student according to their personal interest at the beginning of the course. I also shall suggest specific readings from the primary literature for students that are relevant to their chosen project(s).

I shall assign other readings from the primary literature depending on the weather's effect on fieldwork opportunities, and ask teams of students to lead discussions based on them. Each student will lead several class paper discussions during the summer. At the beginning of the course, I will present a list of potential papers for discussion and in the second week of class, students will vote on which papers interest them most.

Your final exam will consist of a 1-on-1 one hour discussion / interview about core course topics and an oral presentation of your research findings.

Animal Behavior at FLBS is a course that puts you center stage. It calls for you to be extraordinarily active, creatively engaged, and often team-oriented. It invites you to take extreme ownership over your projects, beginning to end. However, I literally am always available to introduce you to research opportunities and support their development. I strive to run the course in the best way possible to help students develop and synergistically fuse conceptual, analytical, and practical evolutionary behavioral ecology skills.

While this course meant to be enormously fun, intellectually stimulating and liberating for mature students, be warned that it is no summer vacation. There will be a small number of 3-day weekends for you to venture out on your own to take recreational trips, e.g., to Glacier National Park, which I highly recommend. But at least as often you probably will need to spend those weekends working on the course at FLBS. I hope the class also will be able to visit some great places during our official time slot.

My phone number is 505-681-3391. You can feel free to call me about any course-related questions. Really


Everyone should have documentation that they are up-to-date on their tetanus immunizations! If you are not sure of your tetnus status get a booster. Rarely, you may need a full series of shots, e.g., if you did not get them as a child. Because of the chance of wounds and cuts during outdoor activities, and the minor risk of being bitten by an animal (e.g., in connection with a major class project, We shall be handling and observing the social behavior of Columbian ground squirrels as a class project; although we have professional quality protective equipment of course, up-to-date tetanus protection is essential.

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