Field Studies in the Evolution of Animal Behavior

Dr. Paul J. Watson - Instructor

An intensive 6-week course to be offered summer 2019 at Flathead Lake Biological Station in the magnificent Northwestern Montana Rockies. Meets Monday through Thursday, June 24th through August 2nd. Go directly to the Biological Station's 2019 summmer session course listings page by clicking here. You can apply online. Do not neglect the scholarship opportunities! Undergraduate or graduate credit is awarded by the University of Montana, Missoula, MT, USA. For University of New Mexico biology students, there may be an option to obtain credits directly from UNM; more on that later.

Behavioral Ecology Rapid Deployment Force, Flathead Lake, Montana, 1994Students of evolution and behavior from across disciplines and the world over are invited to participate; maximum enrollment will be about 12 students. Most of my students are biology majors, but as in years past I will 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 major from American University, Beruit, Lebanon, who immediately went on to do doctoral work in evolutionary antropology at Rutgers.

Aims of the Course
Our big question in this course is, how are animal minds designed to solve niche-specific ecological and ultimately reproductive problems? Behavioral ecology really 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.

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, and which I now teach in an updated form that still honors the approach of my wonderful professor from 1980, the late Dr. Allen Stokes, along with the approach to field work of my superb later mentors (see my main web page). I assisted in teaching rigorous field courses in animal behavior at Cornell University for two terms under Dr. Paul W. Sherman. 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. After a hiatus, and thanks to the recent glorious revivication of the FLBS summer teaching program, we are reinstituting the class in a new 6-week format in 2019.

The course is designed to provide mature, perservering undergraduates, beginning graduates, teachers, and select students from other fields of behavior (e.g., psychology, anthropology, philosophy) 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 take the course seriously 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.

Let me know, by the way, if you wish to communicate with former students.

Behavior Recon Mission, Siyeh Pass, Glacier Park, 2002The 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 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 woud 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.

Siyeh Pass View to the Great PlainsAttentive 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.

Fortunately, most Columbian ground squirrels cannot bite through welder's gloves.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 (40%) 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
All students, especially those with less than junior standing, should directly consult the instructor before enrolling. A strong biological undergraduate background is recommended, but no single course except an evolutionarily oriented introductory biology course for biology major is considered to be a necessary prerequisite. Some of my best students have been mature English and Philosophy majors. But certainly, prior courses in evolution, genetics, statistics, scientific writing, and behavior(!) will enhance most student's experience in this course.

Female fairy moth, Adela brantleyaeCourse 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 to do research. The class needs to be a rapid-deployment-force!

This will be a demanding course for mature dedicated students. Willingness to participate in class activities and independent work 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.

Your instructors will be living at the station and insofar as possible want to be available to you for research support and intellectual discussion during the entire week. We love our work and we are hopelessly afflicted with an overwhelming desire to assist and guide students who wish to pursue theirs!

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:

  1. Watson communes with the mayflies.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)
  2. comparative energetics of female mating resistance in water striders in relation to genitalic morphology
  3. violent male-male competition and chemical communication in a lekking micro-lepidopteran moth (Adela sp.)
  4. female nutritional status, mate number and mate choice in mormon crickets (Anabrus simplex)
  5. aspects of social competition and cooperation among columbian ground squirrels (Spermophilus columbianus)
  6. sex and age-specific reactions to predators in columbian ground squirrels (Spermophilus columbianus)
  7. correlates of territory size and shape in Sorex shrews
  8. fluctuating asymmetry, lek attendance and sexual success in mayflies (Hexigenia sp.)
  9. colony life of Lasius sp. ants, employing large plaster glass-sided ant farms with outdoor access for the ants
  10. fighting ability and copulatory performance (i.e., sexual competitiveness) of male sierra dome spiders (Linyphia litigiosa) in relation to rates of aging
  11. sexual division of labor in nesting birds
  12. the adaptive function of male mimicry by female damselflies (Enallagma boreale)
  13. 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!)
  14. sex pheromones and fighting intensity in sierra dome spiders
  15. nutritional status and sexual receptivity in a water strider (Aquarius remigis)
  16. comparative foraging behavior of various ant species
  17. the energetics of male-male competition in sierra dome spiders
  18. interspecific and intersexual competition and nectar nutritional content in Rufus and Calopie Hummingbirds

For the above mentioned energetics projects, as well as other possible projects that require measurements of energy expenditures in small animals, an oxygen and carbon dioxide respirometry system will be available.

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!

Two of my best-ever students caught GETTING FIELD EXPERIENCE!!!Essays - Several 2-4 page essays focused on evolutionary thought-questions will be assigned during the summer. One on one and group discussion of the essay topics exercises our analytical muscles and help individuals with idiosyncratic problems they have applying evolutionary theory to the adaptationist analysis of animal and human behavior. I am not necessarily interested in a 'correct' answer in these essays. Instead, I look for your having engaged in a logical, evolutionarily informed, and creative process that you have taken time to articulate as well as you can in writing. Essay grades will depend both on content and writing quality.

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.

When you have a semi-final draft of an essay, that is, one whose form and content you are satisfied with, I then would like you to have a fellow student read the essay and give you feedback, not concerning content (that came before, during discussions), but form (i.e., comprehensibility, succinctness, grammar, etc.). Based on the student's comments, try to improve the presentation and then turn in a final draft for instructor evaluation. In other words, I want you to constructively critique one another's writing the way professional writers normally do before they submit something for publication. Note that the goal is not to adopt your peer's writing style, but to hone your own by making it more understandable and grammatically correct. Please get the person who reviewed your semi-final draft to initial the final draft before you hand it in, and please be a conscientious reviewer of the papers of others.

Mating water striders: Gerris incurvatusOral 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 briefly present the results of the project to the class. Presentation of the results of exploratory projects (about 15 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 much of the deeply hallowed and required text, John Alcock's text, "Animal Behavior"in its brand new 11th edition, now co-authored with Dustin R. Rubenstein to reinforce and expand their understanding of topics covered in lecture and discussion. Specific readings from the text will be assigned at the beginning of the course. I also shall suggest specific readings for students that are relevant to their chosen project(s).

I may assign other class readings depending on the weather's effect on fieldwork opportunities, and ask teams of students to lead discussions based on them. Each student may co-lead 1-2 class 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.

Collecting and identifying water striders for study in the Mission Valley, 1998The written exam will cover all major conceptual material covered in lectures. It will have a predominantly short essay format. It will cover textbook and other assigned reading, as well as major discussion topics.

Videos - As a last resort, say, for instance, if more than 2/3rd's of the class have been mauled by a bear, we can watch good Nature videos. We shall discuss the videos with an eye toward evolutionary analysis of the behaviors seen and critique of their scripts. There will also be Neil Young, Warren Zevon, and Stevie Ray Vaughan videos that we'll subject to profound evolutionary analysis.

Behavioral Ecology is a course that puts YOU center stage. It calls for you to be extraordinarily active and creatively engaged - the best way for you to develop and synergistically fuse conceptual, analytical, and practical behavioral ecology skills.

While this course is 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 take recreational trips, e.g., to Glacier National Park, which I recommend, but at least as often you will probably need to spend those weekends working on the course. I hope the class also will be able to visit some great places during some of our official M-Th time slot.
Several times a day I scan my environment for "whiners" and deal with them the good old Montana way... Git away ya dern whiner!!

This page last updated 24 November 2018.