Ecology and Evolutionary Biology

Academic Year 2023 – 2024

General Information

Guyot Hall

Program Offerings:

  • Ph.D.

Director of Graduate Studies:

C. Jessica Metcalf (Fall 2022)

Graduate Program Administrator:


Graduate study in the Department of Ecology and Evolutionary Biology is designed to lead to the degree of Doctor of Philosophy (Ph.D.). 

The special areas of strength in the department are evolution and genomics, ecology and the environment, behavior and organismal biology, conservation biology, and infectious disease. The interests and research of faculty range widely over these areas so that incoming students are able to select their adviser from among several professors working in their chosen discipline. Graduate students also have excellent opportunities for combining several areas for innovative interdisciplinary work.

The graduate program is designed to develop both the breadth and depth of understanding that will enable graduates of the department to respond to future advances in the field. At the same time, students acquire the detailed knowledge and techniques needed to become effective scientists. Each student is guided in developing a comprehensive but flexible course of preparation that is designed to meet their educational needs and goals.


Application deadline
December 1, 11:59 p.m. Eastern Standard Time (This deadline is for applications for enrollment beginning in fall 2024)
Program length
5 years
General Test - optional/not required; Biology Subject Test - optional/not required

Program Offerings

Program Offering: Ph.D.


In consultation with the director of graduate studies, new students work with a graduate committee to develop an individual course work and research program.  During the first two years, members of the committee recommend courses and are available for consultation on planning and initiating research projects.

Though there are few course requirements, all students must enroll in five departmental core courses in their first year. By the end of their second year, students must complete a total of eight core courses, including the field course on tropical ecology, which takes place in January of a student’s first year, and the Responsible Conduct in Research (RCR) course offered every other year. Students may take graduate or undergraduate courses in the department or relevant courses in any other department.  Additional undergraduate courses may be recommended if a student shows an academic deficiency. Graduate students may take additional advanced topic-related courses occasionally throughout their graduate career.

Additional pre-generals requirements

Seminars, Colloquia, and Integrated Research Seminars

Students enjoy the benefits of an excellent series of seminars and colloquia throughout the year, which form an important part of the student’s graduate education and frequently attract faculty and student audiences from several departments such as molecular biology, chemistry, geology, and psychology.  These colloquia feature Princeton faculty, students, and outside speakers and are often given by eminent visiting scientists.  Colloquia are intended to expand the student’s educational experience beyond the bounds of expertise found in the local Princeton community and keep the faculty and students abreast of the latest developments in their field.  Colloquia are held weekly and students are expected to attend colloquia related to their interests.

Research and Thesis Adviser

New students are encouraged to begin research projects as soon as possible.  New students work with the DGS to identify a temporary research adviser and research project.   In some cases, the initial project becomes the thesis topic, but many students work on several smaller projects before beginning the subject of their dissertation. Graduate students are expected to continue their research and training during the summer at Princeton, in the field, or at another laboratory.

Generally, by the end of the second semester, and certainly by the end of the first full year, each student must select a permanent dissertation adviser who will supervise work on the dissertation research.

General exam

The general examination is normally taken in the spring of the second year of study.  However, students can stand for the general examination as early as the end of the first year, provided they have met the Graduate School’s residency requirement. 

The general examination consists of an oral examination, which typically lasts about three hours.  The oral examination is conducted by the student’s dissertation committee, which is chosen in advance by the student and the adviser and is normally composed of four or five Princeton faculty members.  The committee may also include members of other departments within the University. Faculty members from other institutions with special competencies are invited to serve on the dissertation committee should the student’s area of study warrant it.

The student submits a written review of background information relevant to the thesis topic as well as a thesis proposal detailing research objectives, preliminary progress, and future plans. Questions asked during the oral examination focus on the thesis topic and cover all scientific areas relevant to the thesis topic.

Qualifying for the M.A.

The Master of Arts (M.A.) degree is normally an incidental degree on the way to full Ph.D. candidacy and is earned after a student successfully passes the general examination. It may also be awarded to students who, for various reasons, leave the Ph.D. program, provided that this requirement has been met.


All graduate students are required to teach at least two terms during the first four years of the program.  Students who are not supported by an external fellowship, (e.g., NSF, NSERC, etc.) must teach an additional two terms.

Post-Generals requirements

Yearly Committee Meetings

After the general examination, the dissertation committee continues to meet with the graduate student at least once a year to discuss the student's research.  Students are expected to prepare a short written summary of their work before each of these meetings.

Dissertation and FPO

In order to complete the Ph.D., a student consults with the committee members who evaluate the student's dissertation.  Once the committee approves the dissertation, the candidate has two weeks to prepare and present to the department a defense of the work in the final public oral examination for the doctoral degree.

The Ph.D. is awarded after the candidate’s doctoral dissertation has been accepted and the final public oral examination sustained.


  • Chair

    • Jonathan M. Levine
  • Director of Graduate Studies

    • C. Jessica E. Metcalf
  • Director of Undergraduate Studies

    • Robert M. Pringle
  • Professor

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

    • Lindy McBride
    • 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

For a full list of faculty members and fellows please visit the department or program website.

Permanent Courses

Courses listed below are graduate-level courses that have been approved by the program’s faculty as well as the Curriculum Subcommittee of the Faculty Committee on the Graduate School as permanent course offerings. Permanent courses may be offered by the department or program on an ongoing basis, depending on curricular needs, scheduling requirements, and student interest. Not listed below are undergraduate courses and one-time-only graduate courses, which may be found for a specific term through the Registrar’s website. Also not listed are graduate-level independent reading and research courses, which may be approved by the Graduate School for individual students.

EEB 502 - Fundamental Concepts in Ecology, Evolution, and Behavior I

Advanced discussions of ecology, evolution, and behavior focus on 50 fundamental papers. Ecological topics include dynamics and structure of populations, communities, ecosystems, and conservation biology. Behavioral topics include instinct and learning, social behavior, physiological ecology, and the evolution of behavior. Evolutionary topics include speciation, evolutionary and quantitative genetics, molecular evolution, evolutionary stable strategies, and evolution of development. (These are core courses)

EEB 504 - Fundamental Concepts in Ecology, Evolution, and Behavior II

Advanced discussions of ecology, evolution, and behavior focus on 50 fundamental papers. This course continues the discussion of EEB 502, q.v. The two courses are offered in alternate years, and may be taken in either order. Two meetings per week early in the term; one later. (This is a core course.)

EEB 506 - Responsible Conduct in Research (Half-Term)

This course will cover the essential topics of what constitutes responsible conduct in research.

EEB 507 - Recent Research in Population Biology

Systematic reviews of recent literature in areas of ecology, evolution, and animal behavior are made. The general survey of literature is supplemented with detailed discussion of selected research papers of unusual importance and significance. (This is a core course.)

EEB 521 - Tropical Ecology

Intensive three-week field course undertaken in January or February in a suitable tropical locality. There are readings, discussions, and individual projects. The content and location are varied to suit the needs of the participants. Students provide their own travel funds. (This is a core course.)

EEB 522 - Colloquium on the Biology of Populations

Discussion of the central problems of population biology and approaches that have proved fruitful. Topics ranging throughout ecology, evolution, biogeography, and population genetics are usually related to presentations by visiting speakers and students.

EEB 528 - Topics in Conservation (Half-Term)

This half-term topics course offers students exposure to current issues in conservation across the globe. Faculty choose compelling and topical issues that introduce students to the "big" questions in the area, the salient issues under contention, the techniques used to answer the questions, and appropriate case studies that illustrate successes and failures in conservation action.

EEB 531 - Advanced Vertebrate Biology

Topics vary from year to year depending on the instructor's and the students' special interests and may include patterns of reproduction relating to breeding seasons, delayed births, parental care, mating, ovulatory cycles, age of sexual maturity, and viviparity.

EEB 533 - Topics in Ecology

Discussion of the growing points in population and community ecology is held. Topics vary from year to year and include such subjects as the regulation of population numbers, the organization of communities, conservation, and the variation in ecological processes at different spatial and temporal scales.

QCB 515 - Method and Logic in Quantitative Biology (also CHM 517/EEB 517/MOL 515/PHY 570)

Close reading of published papers illustrating the principles, achievements and difficulties that lie at the interface of theory and experiment in biology. Two important papers, read in advance by all students, will be considered each week; emphasis will be on student discussion, not formal lectures. Topics include: cooperativity, robust adaptation, kinetic proofreading, sequence analysis, clustering, phylogenetics, analysis of fluctuations, maximum likelihood methods.

SPI 586B - Topics in STEP (also EEB 516)

These are courses intended to help students develop and apply skills in the application of scientific, technological, and environmental analyses to problems of policy interest. Fall courses are numbered 585, Spring courses are numbered 586.