Ecology and Evolutionary Biology

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

Paul Turner

Director of Graduate Studies
David Vasseur

Professors Richard Bribiescas (Anthropology), Nicholas Christakis (Sociology), Michael Donoghue, Alison Galvani (Public Health), Vivian Irish (Molecular, Cellular & Developmental Biology), Thomas Near, David Post, Jeffrey Powell, Richard Prum, Eric Sargis (Anthropology), Oswald Schmitz (Forestry & Environmental Studies), David Skelly (Forestry & Environmental Studies), Stephen Stearns, Paul Turner, J. Rimas Vaisnys (Electrical Engineering), Günter Wagner

Associate Professors Forrest Crawford (Public Health), Walter Jetz, James Noonan (Genetics), Jeffrey Townsend (Public Health), David Vasseur

Assistant Professors Liza Comita (Forestry & Environmental Studies), Alvaro Sanchez, Carla Staver

Senior Lecturer Marta Martínez Wells

Lecturers Adalgisa Caccone, Linda Puth

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, phylogeny, molecular population genetics, developmental evolution, and evolutionary theory.

Special Admissions Requirements

Applicants should have had training in one of the following fields: biology, mathematics, chemistry, physics, statistics, and/or geology. Candidates are selected, regardless of their major, based on overall preparation for a career in research in ecology and evolutionary biology. Some, planning for careers in applied fields, may have prepared with courses in public policy, economics, and agriculture.

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. All first-year students carry out two research rotations. Students have the option of a rotation over their first summer. Students must participate in (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; and (4) symposia of faculty and graduate student research. In addition, during their first two years of study, graduate students must enroll in a minimum of three additional graduate-level courses (numbered 500 and above); a grade of H must be earned in two of these. Teaching experience is regarded as an integral part of the graduate training program. All students are required to teach three courses, normally at a level 20, typically during their first two years of study.

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 Center for Science and Social Science Information.

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.) Students must pass ten graduate-level courses. At least four courses must be taken for a grade, and students must earn Honors in two courses and maintain an overall average of High Pass. Required courses are: E&EB 500, Advanced Topics in Ecology and Evolutionary Biology; 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. A minimum of five additional graduate-level courses (four taken for a grade) are required.

Additional information on the department, faculty, courses, and facilities is available from Deanna Brunson, Office of the Director of Graduate Studies, Department of Ecology and Evolutionary Biology, Yale University, PO Box 208106, New Haven CT 06520-8106; e-mail,; tel., 203.432.3837; fax, 203.432.2374; website,


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

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 and Walter Jetz

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 515a, Conservation BiologyLinda Puth

An introduction to ecological and evolutionary principles underpinning efforts to conserve Earth's biodiversity. Efforts to halt the rapid increase in disappearance of both plants and animals. Discussion of sociological and economic issues.
MW 10:30am-11:20am

E&EB 520a, General EcologyDavid Vasseur

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.
MWF 10:30am-11:20am

E&EB 523b, Laboratory for Principles of Evolution, Ecology, and BehaviorMarta Wells

Experimental approaches to organismal and population biology, including study of the diversity of life.
MWTh 1:30pm-4:30pm

E&EB 525b, Evolutionary BiologyJeffrey Powell and Alvaro Sanchez

An overview of evolutionary biology as the discipline uniting all of the life sciences. Evolution explains the origin of life and Earth's biodiversity, and how organisms acquire adaptations that improve survival and reproduction. This course uses reading and discussion of scientific papers to emphasize that evolutionary biology is a dynamic science, involving active research to better understand the mysteries of life. We discuss principles of population genetics, paleontology, and systematics; application of evolutionary thinking in disciplines such as developmental biology, ecology, microbiology, molecular biology, and human medicine.
TTh 10:30am-11:20am

E&EB 545b, Responsible Conduct of ResearchDavid Post

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
M 2:30pm-4:30pm

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 575b, Biological OceanographyMary Beth Decker

Exploration of a range of coastal and pelagic ecosystems. Relationships between biological systems and the physical processes that control the movements of water and productivity of marine systems. Anthropogenic impacts on oceans, such as the effects of fishing and climate change. Includes three Friday field trips.
TTh 11:35am-12:50pm

E&EB 620a, Evolutionary and Ecological GeneticsDavid Vasseur

Topics related to analyzing molecular genetic data to answer questions in evolution and ecology. Methods to detect selection in DNA sequences and other molecular data, and landscape genetics, overlaying genetic data on ecological maps from global imaging. Other topics will be determined by interests of participants.
Th 4pm-6pm

E&EB 624b, Advanced EcologyDavid Vasseur

An advanced treatment of ecology, including species interactions, species coexistence theory, species-environment interactions, the maintenance and consequences of biological diversity, spatial ecology, food webs, and eco-evolutionary interactions.
TTh 1pm-2:15pm

E&EB 636b / SOCY 636b, Biosocial ScienceNicholas Christakis

This seminar (with limited enrollment, but open to anyone) covers topics at the intersection of the natural and social sciences, including behavior genetics, gene-environment interactions, social epigenetics, and diverse other topics.
W 4:30pm-6:30pm

E&EB 672b, OrnithologyRichard Prum

An overview of avian biology and evolution, including the structure, function, behavior, and diversity of birds. The evolutionary origin of birds, avian phylogeny, anatomy, physiology, neurobiology, breeding systems, and biogeography.
MWF 9:25am-10:15am

E&EB 673b, Laboratory for OrnithologyRichard Prum

Laboratory and field studies of avian morphology, diversity, phylogeny, classification, identification, and behavior. Must be taken concurrently with E&EB 672.
T 1:30pm-4:30pm

E&EB 680b, Life History EvolutionStephen Stearns

Life history evolution studies how the phenotypic traits directly involved in reproductive success are shaped by evolution to solve ecological problems. The intimate interplay between evolution and ecology.
TTh 11:35am-12:50pm

E&EB 705b, Plant EcologyAnn Staver

Plant ecology is the study of plant interactions with their environment, at the level of individuals, and of how plant-plant interactions mediate environmental interactions at the level of populations, communities, and ecosystems. The course incorporates empirical and theoretical perspectives, emphasizing the empirical origins of concepts in plant ecology and effective empirical tests of conceptual and mathematical predictions. Students read the primary scientific literature extensively, both for content and to build familiarity with methodological standards and the scientific writing.
MW 11:35am-12:50pm

E&EB 713b, Spatial and Environmental Data Analysis in Conservation and Biodiversity ScienceWalter Jetz

The course provides an introduction and hands-on exposure to computational and statistical approaches for the analysis of biodiversity data in a geographical, environmental, and conservation context. After a general overview of relevant hot topics and questions in conservation and ecology and their associated methodologies and data sources, we introduce a set of example questions that we then address with a variety of datasets and methods. A particular focus is the analysis of species distributions and abundances in changing landscapes using remotely sensed environmental information. Beyond broadly available data and methods, students explore new biodiversity-relevant remote-sensing products under development with NASA and prototype tools available through the Yale-based Map of Life project and its partnership with the Google Earth Engine team. Participants gain hands-on experience in spatial analysis and modeling relevant for biodiversity and conservation science and learn about key associated concepts and potential pitfalls. Case studies from forestry, species distribution modeling, biodiversity, and remote sensing data processing. The course meets weekly for 2–3 hours, day and time to be determined. The first organizational meeting takes place on January 20 at 2 pm in OML 201; if you are interested in the course but unable to attend the organizational meeting, please contact the instructor at Prerequisite: open to advanced undergraduates and graduate students (postdocs also welcome) with an interest in advancing their data analysis and modeling skill set and at least some experience in GIS and statistical analysis in R (or willingness to acquire it).

E&EB 725a, Scientific Writing for Ecology and Evolutionary BiologyAnn Staver

This course provides guidance and practice for graduate students in grant and manuscript writing in the fields of ecology and evolutionary biology. Students produce one grant application (NSF GRFP/DDIG or similar) and one manuscript for publication (on a topic of their choice, to contribute to their thesis or other ongoing work).
M 6pm-8pm

E&EB 729a, Microbial Ecology and EvolutionAlvaro Sanchez

This 1-credit graduate seminar course examines various topics in the ecology and evolution of microbes, with an emphasis on prokaryotes (Eubacteria, Archaea) and viruses. Microbiology is an incredibly rich field, where microorganisms are studied from perspectives including the gene, genome, individual, population, community, and ecosystem levels. The course will use discussions of classic and contemporary journal articles to show how species interactions including competition, predation, parasitism, mutualism and microbial communication influence these various levels of biological organization in particular, and evolutionary ecology of microbes in general. Sex and reproduction, genome architecture and reduction, novel evolutionary mechanisms, and challenges imposed by environmental change will be examined from a microbial perspective. The result is an understanding of microbes in their natural habitats ¿whether air, soil, aquatic or host environments ¿ and of the power in using microbes to elucidate fundamental principles in ecology and evolution. Different discussion topics are emphasized in different years, and this year¿s course focuses on the following three areas: studying microbial adaptation in laboratory and natural microcosms, evolutionary ecology of emerging infectious diseases, and the role of microbes in biogeochemical cycling especially in the oceans.
W 1pm-2:50pm

E&EB 810b, Dynamics of Evolving SystemsJ. Rimas Vaišnys

An introduction to the ways evolving biological systems can be described, modeled, and analyzed by using a dynamical systems approach. To use currently fashionable terminology, we develop an individual-based model of the behavior of biological populations, which leads to evolution as an emergent property. In this approach it is possible to construct populations of varying individuals, which can then be combined into larger assemblages, and to modify both the overall environment and the environments at the lower levels, so that aspects often neglected in modeling evolution can be explored and related to any available observational data. Extensive use of the software package Mathematica, but prior experience with the program is not required.
TTh 9am-10:15am

E&EB 842b / ANTH 835b, Primate Diversity and EvolutionEric Sargis

The diversity and evolutionary history of living and extinct primates. Focus on major controversies in primate systematics and evolution, including the origins and relationships of several groups. Consideration of both morphological and molecular studies. Morphological diversity and adaptations explored through museum specimens and fossil casts.
W 1:30pm-3:20pm

E&EB 900a or b, First-Year Introduction to Research and RotationsDavid Vasseur


E&EB 901a, Research Rotation IDavid Vasseur


E&EB 902b, Research Rotation IIDavid Vasseur


E&EB 903b, Independent StudyDavid Vasseur

By arrangement with faculty. Approval of DGS required.

E&EB 930a or b / G&G 703a or b, Seminar in SystematicsJacques Gauthier

A seminar on using molecular evolutionary models in Bayesian phylogenetic analyses. Topics are chosen by the participants but may include "models" in phylogenetics, understanding and comparison of model selection criteria, effects of model under- and overparameterization on parameter value estimates and phylogenetic inferences, and accommodating model uncertainty and model-averaging.

E&EB 950a, Second-Year ResearchDavid Vasseur

By arrangement with faculty.