Department of Ecology and Evolutionary Biology



  • Jonathan M. Levine

Director of Undergraduate Studies

  • Robert M. Pringle

Director of Graduate Studies

  • C. Jessica E. Metcalf


  • Andy P. Dobson
  • Andrea L. Graham
  • Bryan T. Grenfell
  • Lars O. Hedin
  • Simon A. Levin
  • Jonathan M. Levine
  • Stephen W. Pacala
  • Robert M. Pringle
  • Daniel I. Rubenstein
  • Corina E. Tarnita
  • David S. Wilcove

Associate Professor

  • C. Jessica E. Metcalf
  • Christina P. Riehl
  • Mary C. Stoddard
  • Bridgett M. vonHoldt

Assistant Professor

  • Julien F. Ayroles
  • Shane C. Campbell-Staton
  • Sarah D. Kocher
  • Lindy McBride
For a full list of faculty members and fellows please visit the department or program website.

Program Information

The Department of Ecology and Evolutionary Biology (EEB) emphasizes research and teaching from an evolutionary perspective and combines theoretical and empirical approaches. Many of the research projects and courses are interdisciplinary, linking areas that are often treated as separate disciplines. A description of the core areas, faculty interests, and unique features of the program, including details about the department's field programs, can be found on the department's website. Students with an interest in whole-organism and large-scale biological processes—evolution, physiology, disease, behavior, neuroscience, ecology, ecosystem biology, conservation, and climate change—should consider enrolling in EEB.


The Department of Ecology and Evolutionary Biology requires the following prerequisites, which should normally be completed by the end of sophomore year. Prerequisite courses may not be taken using the Pass/D/Fail grading option.

  • Two terms of introductory biology: EEB 211 and either EEB/MOL 214 or EEB/MOL 215)
  • Mastery of calculus to the level of MAT 103 or above, or advanced placement (an AP score of 5 on the AB test or an AP score of 4 on the BC test)
  • Two terms of introductory chemistry (or equivalent, or an AP score of 5)
  • The first term of introductory physics (or the equivalent, or an AP score of 5)
  • A statistics course. SML 201 is preferred, but the following courses will fulfill the requirement: ORF 245, POL 345, PSY 251, ECO 202, SPI 200, SPI 332. It is best to fulfill the statistics requirement before junior year, but it must be fulfilled by the end of the fall term senior year.

Advanced Placement

Advanced placement will be granted to students who received a 5 on the Biology AP exam. Nevertheless, all students planning to major in EEB must take EEB 211 during their first year or sophomore year.

Information and Departmental Plan of Study

Students must successfully complete a minimum of eight upper-level departmental courses, at least six of which must be EEB courses (up to two courses may be offered by other departments). See the department website for a full list of courses.

All EEB concentrators must take EEB 309 and EEB 321 (students are encouraged to take these courses in the fall of their junior year or earlier) as well as one course in two of the following areas:

Behavior and Organismal Biology:  EEB 313; EEB 314; EEB 329; EEB 403; *EEB 404; *EEB 406

Disease Ecology:  EEB 304; EEB 327; EEB 328; EEB 351

Conservation Biology:  EEB 308; *EEB 380; EEB 417

Mathematical and Computational Biology:  EEB 324; EEB 325

* offered as part of semester abroad program (described below)

For the remaining departmentals, students may select any EEB course or those offered by other departments as follows. The first semester of organic chemistry and biochemistry (MOL 345) can each be counted as a departmental course (the second semester of organic chemistry will not count). Other MOL courses might count with preapproval from the director of undergraduate studies (DUS) via the undergraduate program administrator (UPA).  With preapproval from the DUS via the UPA, students may take one science-relevant course on policy or society. 

Only Princeton courses can count as departmentals; the one exception is for preapproved courses taken during a study-abroad term, but the letter grades will not transfer. No course taken Pass/D/Fail can be counted as a departmental or to satisfy requirements of the major. The minimum grade for a course to count as a departmental is C-. 

Integrated Biology Sequence: Students who have completed the two-year sequence need to meet with the EEB director of undergraduate studies to discuss placement in upper-level EEB courses.

Pre-medical Students:

Students interested in medical school should consider taking EEB 314 Comparative Physiology, which will count toward one of the four required areas of study. In addition, medical schools require two terms of physics; physics can be delayed to senior year if necessary. The Health Profession advisers recommend that students who wish to study abroad, especially in EEB’s field semesters, should do so. They recommend contacting them early to formulate a plan of study that allows students to fulfill the premedical prerequisites and allows them to study abroad during spring of their junior year.

    Early Concentration

    Students who are passionate about solving problems in the areas of ecology, evolution, behavior, conservation, and disease may apply for early concentration in ecology and evolutionary biology. Students who have advanced placement in biology and have taken at least one upper-level EEB course, in addition to having completed many of the department's required pre- and co-requisite courses with grades of at least B+, are eligible. Early concentrators will begin independent work during the spring of their sophomore year. Students interested in early concentration should contact a director of undergraduate studies at the end of the first year or at the start of sophomore year.

    Independent Work

    Junior Independent Work:  Early in the fall semester, students will attend periodic meetings on Tuesday evenings that will include presentations by EEB faculty members on their research and on opportunities for senior independent work. After the faculty presentations, which will take place early in the semester, students will identify those faculty with whom they are interested in working. If the faculty member agrees to take on a particular student, together they will develop a plan of study.

    During the second half of the fall term, students participate in a tutorial in which they work closely with a faculty member to address a problem and write a first junior paper. In the spring semester, students carry out a program of independent research with their faculty adviser, which can include empirical or theoretical work. Students will summarize this research project in a second junior paper, which can be structured either as a research project, a literature review, or a research proposal. The second junior paper is due in early May.

    Senior Independent Work: During the fall of junior year, each student selects a senior thesis adviser (see above). The adviser and the student choose a research project that the student generally pursues during the summer preceding senior year and both terms of the senior year. The research project can involve primarily laboratory, field, data mining, theoretical, or library study that will be written and presented as a senior thesis.

    Senior Departmental Examination

     A one-hour oral examination, consisting of a defense of the thesis research and general questions in the biological sciences, will be held in May.

    Study Abroad

    Special Features of the Plan of Study. EEB offers two tropical field study programs: a semester in Panama, and a semester in Kenya. Four courses, built around experiential learning, are taken in sequence at each location typically during the spring of junior year. Please note that seniors who are making good progress with their thesis are encouraged to consider fulfilling their last two courses in the spring of senior year by attending the first half of the semester in Kenya or Panama. Seniors who choose to do this would return to campus at spring break to complete the senior thesis. Details of these programs can be found on the department's website.

    Princeton’s Tropical Field Programs. Students interested in learning about or undertaking research in the tropics have a number of options.

    1. Panama. The department offers a spring term in Panama in conjunction with the Smithsonian Tropical Research Institute. Students take four intensive three-week courses in sequence, beginning with an introduction of key concepts in tropical ecology and conservation. The program also includes courses on coral reefs, parasitology, and anthropology (focusing on Pre-Columbian peoples and their land-use practices).

    2. Kenya. This four-course program on Tropical Biology and Sustainability, also taught in three-week segments, takes place at Princeton University's Mpala Research Centre in central Kenya and other sites in Kenya, in collaboration with scientists there, EEB faculty, other appropriate faculty from Princeton University, and faculty from Columbia University. Columbia students also participate in the program. The courses delve into the ecology of savannas, conservation in Africa, the natural history of mammals, tropical agriculture, engineering, and field hydrology and paleoecology.

    3. Other. Individual students are welcome to pursue other independent field opportunities, with scientists from the Smithsonian Institution and the Bermuda Institute of Ocean Sciences, or other research institutions, such as the School for Field Studies at the University of Cape Town in South Africa, or James Cook University in Australia.

    Additional Information

    Program in Teacher Preparation. As the need for qualified science teachers increases, some students may wish to earn a teaching certificate. Working with the director of undergraduate studies and the teacher preparation program, an appropriate course of study can be arranged.


    EEB 211 Life on Earth: Mechanisms of Change in Nature Fall SEL

    An examination of how life evolved and how organisms interact to shape the natural world. Why did the dinosaurs disappear? What mechanisms can produce the chameleon's camouflage or the giraffe's long neck? Why do ecosystems contain a wide diversity of species when competition between them should eliminate all but a few? How will life on earth change with increasing human domination of the planet? These and other questions related to the origin and future of life, conflict and cooperation between species, and dynamics of entire ecosystems will be explored. This course is required for EEB majors and fulfills a requirement for medical school. Instructed by: J. Levine, S. Kocher

    EEB 214 Introduction to Cellular and Molecular Biology (See MOL 214)

    EEB 304 Disease Ecology, Economics, and Policy (See ENV 304)

    EEB 308 Conservation Biology (also
    ENV 365
    ) Fall SEN

    Students will use ecological principles and policy analysis to examine conflicts between human activities such as farming, forestry, and infrastructure development, and the conservation of species and ecosystem services. Two lectures, one preceptorial. Instructed by: D. Wilcove

    EEB 309 Evolutionary Biology Fall

    All life on Earth has evolved and continues to evolve. This course will explore evolution at both the molecular and organismal level. We will examine the features that are universal to all life and that document its descent from a common ancestor that lived over 3 billion years ago. Topics include the origin of life, the evidence for natural selection, methods for reconstructing evolutionary history using DNA, population genetics, genome evolution, speciation, extinction, and human origins. This course will provide you with the basic tools to understand how evolution works and can produce the incredible diversity of life on our planet. Instructed by: J. Ayroles

    EEB 311A Animal Behavior Not offered this year SEN

    An examination of the mechanisms and evolution of the behavior of humans and other animals. Topics include the sensory worlds of animals, the nature of instinct, neural mechanisms of perception, comparative studies of communication, learning, cognition, mate choice, and social behavior, and the biology of human development and language acquisition. Two 90-minute lectures, one preceptorial. Instructed by: J. Gould

    EEB 314 Comparative Physiology Spring

    The study of how animals function with emphasis on the integration of physiological processes at the cellular, organ, and whole organism levels in ecological and evolutionary contexts. Comparisons among species and higher taxa are used to illustrate general physiological principles and their evolutionary correlates. Three lectures, one three-hour laboratory. Prerequisite: 210 or 211. Instructed by: Staff

    EEB 315 Human Adaptation (See ANT 215)

    EEB 321 Ecology: Species Interactions, Biodiversity and Society (also
    ENV 384
    ) Fall SEL

    How do wild organisms interact with each other, their physical environments, and human societies? Lectures will examine a series of fundamental topics in ecology -- herbivory, predation, competition, mutualism, species invasions, biogeographic patterns, extinction, climate change, and conservation, among others--through the lens of case studies drawn from all over the world. Readings will provide background information necessary to contextualize these case studies and clarify the linkages between them. Precepts and fieldwork will explore the process of translating observations and data into an understanding of how the natural world works. Instructed by: R. Pringle

    EEB 324 Theoretical Ecology Spring QCR

    Current and classical theoretical issues in ecology and evolutionary biology. Emphasis will be on theories and concepts and on mathematical approaches. Topics will include population and community ecology, epidemiology and evolutionary theory. Two lectures, one preceptorial/computer laboratory. Prerequisite: one year of calculus. Instructed by: S. Levin

    EEB 325 Mathematical Modeling in Biology and Medicine Fall

    How can mathematical modeling help to illuminate biological processes? This course examines major topics in biology through the lens of mathematics, focusing on the role of models in scientific discovery. Students will learn how to build and analyze models using a variety of mathematical tools. Particular emphasis will be placed on evolutionary game theory. Specific topics will include: the evolution of cooperation and of social behavior from bacteria to humans; the evolution of multicellularity; the somatic evolution of cancer; virus dynamics (within host and within populations); and multispecies interactions and the evolution of mutualisms. Instructed by: C. Tarnita

    EEB 327 Immune Systems: From Molecules to Populations (also
    MOL 327
    GHP 327
    ) Fall SEN

    Why is there immunological polymorphism in animal populations? Why do immune systems work as they do? This course examines the theories of host-parasite coevolution, including optimal host resource allocation to immune defense in light of parasite counter-strategies, and assesses the empirical evidence by which these theories are tested. Students look at the evolutionary ecology of mechanisms used by immune systems to recognize and kill parasites, finding similarities across animal taxa. Finally, students will map immune mechanisms onto host phylogenies to understand the order in which different mechanisms arose over evolutionary time. Instructed by: A. Graham

    EEB 328 Ecology and Epidemiology of Parasites and Infectious Diseases (also
    GHP 328
    ) Spring SEL

    An introduction to the biology of viruses, bacteria, fungi, protozoa, worms, arthropods, and plants that are parasitic upon other animal and plant species. The major emphasis will be on the parasites of animals and plants, with further study of the epidemiology of infectious diseases in human populations. Studies of AIDS, anthrax, and worms, and their role in human history, will be complemented by ecological and evolutionary studies of mistletoe, measles, myxomatosis, and communities of parasitic helminths. Limited to students in the Tropical Ecology Program in Panama. Instructed by: A. Dobson

    EEB 332 Pre-Columbian Peoples of Tropical America and Their Environments (also
    LAS 350
    ) Spring SA

    An intensive course on the pre-European history of Amerind cultures and their environments in the New World tropics. Topics include the people of tropical America; development of hunting/gathering and agricultural economies; neotropical climate and vegetation history; and the art, symbolism, and social organization of native Americans. Daily lectures, field trips, and laboratory experiences and incorporates methods and problems in field archaeology, paleoethnobotany and paleoecology, and archaeozoology. Limited to students in the Tropical Ecology Program in Panama. This course does not count as an EEB departmental. Prerequisite: EEB 321. Instructed by: Staff

    EEB 336 The Diversity of Brains (See PSY 336)

    EEB 338 Tropical Biology (also
    LAS 351
    ) Spring SEL

    This intensive field course, at various sites in Panama, examines the origins, maintenance, and major interactions among elements of the tropical-terrestrial biota. Study topics include identification of common orders and families of neotropical organisms; tropical climate and hydrology; biotic interactions; and contemporary and historical factors in shaping tropical landscapes, with emphasis on the Isthmian Landbridge and subsequent floral and faunal interactions. Two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. Instructed by: Staff

    EEB 346 Biology of Coral Reefs Spring SEL

    This intensive field course provides an in-depth introduction to the biology of tropical coral reefs, with an emphasis on reef fish ecology and behavior. Students learn to identify fishes, corals, and invertebrates, and learn a variety of field methods including underwater censusing, mapping, videotaping, and the recording of inter-individual interactions. Two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Snorkeling in open ocean and walking in wild terrain is common. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. Instructed by: Staff

    EEB 350 Vertebrate Tropical Ecology Spring

    This intensive field course addresses the life-history characteristics of tropical vertebrates and the physiological traits that underlie them. Students will learn how tropical life histories differ from those in the temperate zone and will use eco-physiological techniques while conducting experiments and observations at a Smithsonian Institute field station. Two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. Instructed by: Staff

    EEB 351 Epidemiology: An Ecological and Evolutionary Perspective (See GHP 351)

    EEB 404 Natural History of Mammals Not offered this year SEL

    Students examine how mammals interact with diverse and potentially conflicting features of their environment in order to understand the concepts, methods, and material of comparative natural history. Perspectives include morphology, identification, evolution, ecology, behavior, habitat, and conservation. Original observations and experiments culminate in class, group, and individual research projects. This intensive field course entails two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Limited to students in the Tropical Ecology Program in Kenya. Prerequisite: 211 and 321. Instructed by: D. Rubenstein

    EEB 417A Ecosystems and Global Change (also
    ENV 417A
    ) Not offered this year SEN

    An introduction to the concepts, approaches, and methods for studying complex ecological systems, from local to global scales. Students will examine nutrient cycling, energy flow, and evolutionary processes, with emphasis on experimental approaches and comparisons between terrestrial, freshwater, and marine ecosystems. Particular attention will be on effects of human activities, including climate change, biodiversity loss, eutrophication, and acid rain. Prerequisites: 210 or 211 or equivalent; CHM 301 or equivalent. Two 90-minute classes. Instructed by: L. Hedin

    EEB 417B Ecosystems and Global Change (also
    ENV 417B
    ) Not offered this year SEL

    An introduction to the concepts, approaches, and methods for studying complex ecological systems, from local to global scales. Students will examine nutrient cycling, energy flow, and evolutionary processes, with emphasis on experimental approaches and comparisons between terrestrial, freshwater, and marine ecosystems. Particular attention will be on effects of human activities, including climate change, biodiversity loss, eutrophication, and acid rain. Prerequisites: 210 or 211 or equivalent; CHM 301 or equivalent. Two 90-minute classes, one three-hour laboratory. Instructed by: L. Hedin

    EEB 419 Environmental Microbiology (See GEO 417)