Ecology and Evolutionary Biology

Osborn Memorial Laboratories, 203.432.3837
M.S., Ph.D.

Thomas Near

Director of Graduate Studies
Martha Muñoz

Professors Richard Bribiescas (Anthropology), Craig Brodersen (School of the Environment), Nicholas Christakis (Sociology), Liza Comita (School of the Environment), Casey Dunn, Erika Edwards, Vanessa Ezenwa, Vivian Irish (Molecular, Cellular, & Developmental Biology), Walter Jetz, Thomas Near, David Post, Jeffrey Powell, Richard Prum, Eric Sargis (Anthropology), Oswald Schmitz (School of the Environment), David Skelly (School of the Environment), Jeffrey Townsend (Public Health), Paul Turner, David Vasseur

Associate Professors Forrest Crawford (Public Health), James Noonan (Genetics), Carla Staver, Alison Sweeney

Assistant Professors Jennifer Coughlan, Nathan Grubaugh (Public Health), Martha Muñoz, C. Brandon Ogbunu, Eric Slessarev, Serena Tucci (Anthropology), Michelle Wong

Senior Lecturer Marta Martínez Wells

Lecturers Adalgisa Caccone, Gordon Geballe, Joshua Moyer, Linda Puth

Research Scientist Mary Beth Decker

Fields of Study

The Department of Ecology and Evolutionary Biology (E&EB) offers training programs in organismal biology, ecology, and evolutionary biology including molecular evolution, phylogenetics, molecular population genetics, developmental evolution, and evolutionary theory.

Special Requirements for the Ph.D. Degree

Each entering student, in consultation with the director of graduate studies (DGS), develops a specific program of courses, seminars, laboratory research, and independent reading tailored to the student’s interests, background, and goals. There are normally no foreign language requirements. The course requirements to advance to candidacy in E&EB are (1) E&EB 500* and E&EB 501*, Advanced Topics in Ecology and Evolutionary Biology; (2) E&EB 545*, a course on the responsible conduct of research; (3) weekly E&EB seminars; (4) symposia of faculty and graduate student research; (5) two research rotations (E&EB 901*, Research Rotation I, and E&EB 902*, Research Rotation II) in the first two years; and (6) a minimum of three additional graduate-level courses (numbered 500 and above) with a grade of Honors (H) in at least two of these.

Teaching experience is regarded as an integral part of the graduate training program. All students are required to teach three courses, typically during their first three years of study. Students who require additional support from the Graduate School must teach additional terms, if needed, after they have fulfilled the academic teaching requirement.

By the middle of the fourth term of study, each student organizes a formal pre­prospectus consultative meeting with the student’s advisory committee to discuss the planned dissertation research. Before the beginning of the fifth term, students present and defend their planned dissertation research at a prospectus meeting, at which the department determines the viability and appropriateness of the student’s Ph.D. proposal. A successful prospectus meeting and completion of course requirements results in admission to candidacy for the Ph.D. The remaining requirements include completion, presentation, and successful defense of the dissertation, and submission of copies of the dissertation to the Graduate School and to the Marx Science and Social Science Library.

In cases where the dissertation committee decides that preliminary field work during the summer after the fourth term is necessary prior to the prospectus, the prospectus meeting can be delayed by one term. A request for a delay must come from the dissertation committee adviser and must be approved by the DGS. In these exceptional cases, admission to candidacy may not be required for registration for the third year of graduate study.

Honors Requirement

Students must meet the Graduate School’s requirement of Honors in two courses by the end of the fourth term of study. The E&EB department also requires an average grade of at least High Pass in course work during the first two years of study.

Master’s Degree

M.S. (en route to the Ph.D.) The course requirements for the M.S. are the same those as for advancing to candidacy in the Ph.D. program: Required courses are: E&EB 500 and E&EB 501, Advanced Topics in Ecology and Evolutionary Biology; E&EB 545, Responsible Conduct of Research; E&EB 901, Research Rotation I; and E&EB 902, Research Rotation II. These courses are taken Sat/Unsat. A minimum of three additional graduate-level, elective courses are required and must be taken for a grade. Students must earn Honors in at least two courses and maintain an overall average of High Pass.

Additional information on the department, faculty, courses, and facilities is available from Kelly Pyers, Registrar, Department of Ecology and Evolutionary Biology, Yale University, PO Box 208106, New Haven CT 06520-8106; email,; tel., 203.432.3837;


E&EB 500a and E&EB 501b, Advanced Topics in Ecology and Evolutionary BiologyStaff

Topics to be announced. Graded Satisfactory/Unsatisfactory.
M 2:30pm-4:30pm

E&EB 510a / S&DS 501a, Introduction to Statistics: Life SciencesJonathan Reuning-Scherer

Statistical and probabilistic analysis of biological problems, presented with a unified foundation in basic statistical theory. Problems are drawn from genetics, ecology, epidemiology, and bioinformatics.
TTh 1pm-2:15pm

E&EB 520a, General EcologyCarla Staver

A broad consideration of the theory and practice of ecology, including the ecology of individuals, population dynamics and regulation, community structure, ecosystem function, and ecological interactions on broad spatial and temporal scales. Topics such as climate change, fisheries management, and infectious disease are placed in an ecological context.
MW 11:35am-12:50pm

E&EB 523Lb, Laboratory for Evolution, Functional Traits, and the Tree of LifeMarta Wells

Experimental approaches to organismal and population biology, including study of the diversity of life.

E&EB 545b, Responsible Conduct of ResearchStaff

This five-week discussion seminar considers issues related to the responsible conduct of research. Topics addressed include research misconduct, plagiarism, data acquisition and management, mentoring and collaboration, authorship and peer review, the use of animals and humans in scientific research, sexual harassment, diversity, and balancing professional and personal life. Graded Satisfactory/Unsatisfactory.  0 Course cr

E&EB 550a, Biology of Terrestrial ArthropodsMarta Wells

Evolutionary history and diversity of terrestrial arthropods (body plan, phylogenetic relations, fossil record); physiology and functional morphology (water relations, thermo-regulation, energetics of flying and singing); reproduction (biology of reproduction, life cycles, metamorphosis, parental care); behavior (migration, communication, mating systems, evolution of sociality); ecology (parasitism, mutualism, predator-prey interactions, competition, plant-insect interactions).
TTh 11:35am-12:50pm

E&EB 551La, Laboratory for Biology of Terrestrial ArthropodsMarta Wells

Comparative anatomy, dissections, identification, and classifications of terrestrial arthropods; specimen collection; field trips.
W 1:30pm-4:30pm

E&EB 555a, InvertebratesCasey Dunn

An overview of animal diversity that explores themes including animal phylogenetics (evolutionary relationships), comparative studies of evolutionary patterns across species, organism structure and function, and the interaction of organisms with their environments. Most animal lineages are marine invertebrates, so marine invertebrates are the focus of most of the course. Concurrent enrollment in E&EB 556L is not required.
TTh 11:35am-12:50pm

E&EB 556La, Laboratory for InvertebratesCasey Dunn

The study of invertebrate anatomy and diversity in a laboratory and field setting. Activities include examination of live animals and museum specimens, as well as local field trips. Some field trips fall on weekends. Must be taken concurrently with E&EB 555.  ½ Course cr
Th 1:30pm-4:30pm

E&EB 620b, Community EcologyDavid Vasseur

This course covers core questions in community ecology related to species interactions, species coexistence theory, species-environment interactions, the consequences of biological diversity, spatial ecology, food webs, and eco-evolutionary interactions. Lectures emphasize the theoretical and conceptual foundations of these topics and incorporate the empirical and experimental evidence supporting and confronting contemporary views.
TTh 1pm-2:15pm

E&EB 622a, Evolutionary GeneticsJennifer Coughlan

Genetic variation is the currency by which natural selection is translated into evolutionary change. In this course we dissect patterns of genetic variation using an evolutionary mindset to ultimately understand what shapes genetic variation in nature and the potential for species to adapt to new and changing environments. This class unites two foundational fields of evolutionary genetics: quantitative genetics (the study of the genetic basis of complex traits) and population genetics (the study of gene variant frequencies across time and space), with an ultimate goal of understanding evolutionary change in nature. Although this course is lecture based, there is much opportunity for hands-on learning. Students use real-life and simulated genetic data to map the genetic basis of traits and investigate the evolutionary forces responsible for shaping genetic variation in nature. We also discuss how quantitative and population genetics theory are applied to the modern genomic era, particularly in the context of detecting genomic signatures of adaptation. Last, we discuss the application of evolutionary genetics to human populations, including the usefulness and missteps of these applications for science and society.
MW 1pm-2:15pm

E&EB 635a, Evolution and MedicineBrandon Ogbunu

Introduction to the ways in which evolutionary science informs medical research and clinical practice. Diseases of civilization and their relation to humans’ evolutionary past; the evolution of human defense mechanisms; antibiotic resistance and virulence in pathogens; cancer as an evolutionary process. Students view course lectures online; class time focuses on discussion of lecture topics and research papers. Prerequisites: BIOL 101BIOL 104.
TTh 1pm-2:15pm

E&EB 712a, Foundations of EcologyDavid Vasseur

This seminar course familiarizes students with foundational concepts and themes in ecology and how they have changed over time. Each week we read and discuss two papers: one classic paper selected from the recently published volume Foundations of Ecology II: Classic Papers with Commentaries (Eds. Miller and Travis, 2022) covering the period 1970–1995, and one related contemporary paper published after 2010. We discuss how the concepts and themes introduced in classic papers have influenced the field of ecology and consider how new tools, data, and insights have advanced, diminished, or changed their impact. The Foundations book covers many topics, arranged into six core areas. Readings cover all six areas, but the included content varies depending on the interests of the class. Students are responsible for choosing one classic paper from Foundations, pairing it with one contemporary paper and leading the discussion during the class meeting. Students also submit short weekly “reflections” in response to a prompt.
Th 9:25am-11:15am

E&EB 721a, Foundations of Terrestrial EcologyMichelle Wong

Intended for graduate students, this seminar course brings a historical perspective to understanding current questions and approaches in terrestrial ecology, ranging from evolutionary, community, landscape, to ecosystem ecology. We read and discuss foundational papers and related current papers, and we identify future directions, opportunities, and challenges for the different sub-fields. The course allows students to critically examine and engage with some scientific work that has laid the findings and concepts that are foundational as they develop conceptual and methodological approaches to their own research. Starting in weeks three or four, each student takes a turn leading discussion and selecting a relevant current paper to that week’s topic. Students write a total of seven précis on the topics of their choosing, with at least two completed by week six.
T 9:25am-11:15am

E&EB 830b, The Ecology of the Great PandemicsBrandon Ogbunu

In this course we examine principles of the ecology of infectious disease in light of three pandemics: the 1918 influenza pandemic, the HIV/AIDS pandemic, and the COVID-19 pandemic. The course covers principles of zooneses, disease emergence, herd immunity, basic vaccinology, and other fundamental concepts. It also focuses on social and cultural factors that fomented these pandemics.
Th 3:30pm-5:20pm

E&EB 856a, Special Topics in the Ecology and Evolution of Infectious DiseasesVanessa Ezenwa

Historically, pathogens and the diseases they cause were viewed largely from a biomedical perspective focused on interactions between pathogens and their human hosts. However, in the last few decades, the importance of studying pathogens from an ecological and evolutionary perspective has gained significant traction. These perspectives inform our understanding of almost all aspects of pathogen-host interactions from transmission dynamics and zoonotic disease spillover to the evolution of virulence and drug resistance. In this seminar, we dissect current and classic literature on the ecology and evolution of infectious diseases. Specifically, we: (i) discuss fundamental concepts in the field; (ii) identify persistent knowledge gaps; and (iii) explore opportunities for linkages between ecological, evolutionary, and biomedical perspectives.
W 1:30pm-3:20pm

E&EB 901a or b, Research Rotation IStaff


E&EB 902a or b, Research Rotation IIStaff


E&EB 930a / EPS 703a, Seminar in SystematicsJacques Gauthier

Topics and class time are chosen by the participants, and have included reading books and/or a series of papers on particular topics (e.g., homology; morphological phylogenetics; evolution of egg colors and exposed nesting in dinosaurs/birds; origin of snake ecology; conflicts between morphology and molecules; role of fossils in phylogenetic inference).