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Department of Ecology and Evolutionary Biology


Lars O. Hedin

Acting Chair

Bryan T. Grenfell (spring)

Departmental Representative

Daniel I. Rubenstein (fall) 

Stephen W. Pacala (spring)

Director of Graduate Studies

Andrew P. Dobson


Andrew P. Dobson

Bryan T. Grenfell, also Woodrow Wilson School

Lars O. Hedin, also Princeton Environmental Institute

Laura F. Landweber

Simon A. Levin

Stephen W. Pacala

Daniel I. Rubenstein

David S. Wilcove, also Woodrow Wilson School

Associate Professor

Peter Andolfatto, also Lewis-Sigler Institute for Integrative Genomics

Andrea L. Graham

Assistant Professor

Julien Ayroles, also Lewis-Sigler Institute for Integrative Genomics

Carolyn S. McBride, also Princeton Neuroscience Institute

C. Jessica E. Metcalf, also Woodrow Wilson School

Robert M. Pringle

Christina P. Riehl

Corina E. Tarnita

Bridgett M. vonHoldt

Associated Faculty

Asif A. Ghazanfar, Psychology, Princeton Neuroscience Institute

Kelly K. Caylor, Civil and Environmental Engineering

Ignacio Rodríguez-Iturbe, Civil and Environmental Engineering

Bess B. Ward, Geosciences

Information and Departmental Plan of Study

Courses in the biological sciences at Princeton are offered in two departments. Students with an interest in whole-organism and large-scale processes--evolution (including molecular evolution and developmental evolution), physiology, disease, behavior, neuroscience, ecology, ecosystem biology, conservation, and climate change--should enroll in the Department of Ecology and Evolutionary Biology (EEB). Those with interests in molecular, cellular, and developmental processes should enroll in the Department of Molecular Biology (MOL). Both departments provide an excellent background for medical school.

The EEB department emphasizes research and teaching from an evolutionary perspective, combining theory and empiricism and linking areas that are often treated as separate disciplines. Many of the research projects and courses are interdisciplinary. A description of these 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.

Every student considering majoring in EEB should attend the departmental open house held in the spring term. It introduces potential majors to departmental courses, faculty, and the wide-ranging research and field-course opportunities open to students. See the department's website for examples of recent student research activities.

Advanced Placement

Advanced placement will be granted to students who received a 5 on the Biology AP exam. Nevertheless, all students planning on majoring in EEB must take EEB 211.


The Department of Ecology and Evolutionary Biology requires the following prerequisites, which should normally be completed by the end of sophomore year. (Two exceptions, physics and statistics, are noted below.)

- Two terms of introductory biology: EEB 211 and either EEB/MOL 214 or EEB/MOL 215);
- Mastery of calculus to the level of MAT 102 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). [Note that medical schools require two terms of physics. Physics can be delayed to the junior or senior year if necessary but preferably should be taken earlier].
- An introductory statistics course: ECO 202, ORF 245, POL 345, PSY 251, SOC 301, or WWS 332 (ORF 245 or PSY 251 are preferred). The statistics requirement must be fulfilled by the end of fall term senior 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 having advanced placement in biology and having 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 departmental representative at the end of the freshman year or at the start of the sophomore year.

Program of Study

Requirements. The following courses are required for graduation:

1. Students in ecology and evolutionary biology must successfully complete at least eight upper-level departmental courses, at least five of which must be EEB or MOL courses. One of the eight must be an EEB lab course; 314, 324, 417B, or MOL 350 and all field courses count as lab courses.

Choice of Courses. Students must take at least one course from each of the following three general areas:

Behavior and Physiology: 301, 311, 314, 350, 404, PSY 315;
Evolution and Genetics: 309, 320, 414;
Ecology and Conservation: 308, 312, 313, 321, 322, 324, 328, 338, 346, 380, 417.

2. Organic chemistry: CHM 301 or 303; (Students interested in mathematical biology may take one of the following courses instead of organic chemistry: PHY 203; MAT 201, 202, 203, 204; COS 126, 226); or two upper-level courses in an aligned area with permission of the departmental representative);

3. Integrated Biology Sequence: Students who have completed the two-year sequence need to meet with the EEB departmental representative to discuss placement in upper-level EEB courses.

Students may take 309, 313, and 321 in the fall term of junior year to sample all three areas. Pay particular attention to the timing of the courses that are taught in alternate years.

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-. Only one course with a policy perspective can be counted as a departmental. Only Princeton courses can count as departmentals; the one exception is for pre-approved courses taken during a study-abroad term.

Special Features of the Plan of Study. EEB offers two tropical field study programs: a semester in Panama, and a semester in Kenya. Details of these programs can be found on the department's website.

Specializations. Students interested in specializing within the department should enroll in the courses listed below.

Theoretical biology: 309, 320, 321, 322, 324, and 325
Field biology: 312, 321, and a tropical field program
Conservation biology: 308, 417, and a tropical field program
Medical biology: 314, 327, and 328
Human biology: 301, 306, 315, PSY 330, and 336
Molecular evolution: 309, 320 or 414, plus other courses in MOL

Note: MOL 214 or 215 is required for medical school. A full year of physics is required for medical school and by some graduate programs (especially in neurobiology). A full year of organic chemistry is required for medical school, and is a prerequisite for many molecular biology courses; both terms count as EEB departmentals. At least one term of organic chemistry is usually required by graduate programs in biology. Completing the organic chemistry requirement before junior year is strongly recommended.

Independent Work

Junior Independent Work. Students attend a weekly junior tutorial in the first half of the fall term. Faculty members present summaries of their work and research opportunities open for senior independent work. During the second half of the fall term, students work closely with one faculty member to investigate a problem using the current literature, and then write a paper. In the spring semester, students carry out a program of independent work with a faculty adviser. In some instances this may include empirical or theoretical work. Either a paper summarizing this project or a research proposal for the senior thesis is due in early May.

Senior Independent Work. During the fall of the junior year each student selects a senior thesis adviser. The adviser and the student choose a research project that the student generally pursues during the summer preceding the 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.

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). Prerequisite: EEB 321 or an equivalent introductory ecology course.

2. Kenya. This four-course program, 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 participate in the program. The courses delve into the ecology of savannas, conservation in Africa, the natural history of mammals, tropical agriculture and engineering and field hydrology. Prerequisite: EEB 321 or an equivalent introductory ecology course.

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.

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


EEB 211 Life on Earth: Chaos and Clockwork of Biological Design (also MOL 211)   Fall STL

An examination of how life evolved and how organisms function. Design--'intelligent' and otherwise--will provide a unifying theme. Why do some microbes produce slime and others do not? Why are males brightly colored in some species, but in others females are the showy sex? Why do humans have knees that fail whereas horses and zebras do not? These and other 'why is it so' questions related to the origin and history of life, genetic code, biochemistry, physiology, morphology and body plans, sex and reproduction, cooperation, and ecosystems will be explored. This course is required of all EEB majors and fulfills a requirement for medical school. S. Pacala, D. Rubenstein

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

EEB 215 Quantitative Principles in Cell and Molecular Biology (see MOL 215)

EEB 255A Life in the Universe (see GEO 255A)

EEB 255B Life in the Universe (see GEO 255B)

EEB 301 Evolution and the Behavior of the Sexes (also GSS 301)   Not offered this year STN

Psychological, biological, and cross-cultural approaches to the study of sex and gender. Topics include biological components and development of sex differences; acquisitions of gender identity; social organization of key life cycle events; evolutionary considerations in the study of sex differences. One 90-minute lecture, one 90-minute class. Staff

EEB 302 Practical Models for Environmental Systems (see ENV 302)

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

EEB 305 Water, Energy, and Ecosystems (see CEE 307)

EEB 306 Human Evolution (see ANT 206B)

EEB 308 Conservation Biology   Fall STN

A detailed application of ecological principles to the conservation of biological resources, including island biogeography, population genetics and viability, and landscape ecology. Analysis of case studies in conservation. Individual project on a conservation issue of the student's choice. Two lectures, one preceptorial. Staff

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. B. vonHoldt

EEB 311A Animal Behavior   Spring STN

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. J. Gould

EEB 312 Marine Biology (also ENV 312)   Not offered this year STL

An intensive four-week course during June in Bermuda. Covers elements of the ecology, evolution, physiology, and behavior of marine organisms and ecosystems. Habitats examined will include the intertidal zone, seagrass beds, mangroves, and the open ocean, with special attention to coral reefs. Topics range from the physiology and behavior of individuals in the habitat, to the flow of energy, predator/prey interactions, symbioses, and population dynamics. Prerequisites: 210 or 211, ability to swim. Three hours of lectures, three hours of laboratory and field trips per day. 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. Staff

EEB 315 Human Adaptation (see ANT 215)

EEB 320 Molecular Evolution (also MOL 330)   Spring

How and where did life evolve? This advanced seminar will discuss the evolution of the molecules that sustain life (DNA, RNA and proteins) at both the micro and macro evolutionary levels. We will explore the role of these molecules in the origin and continued evolution of life. Topics include the origin of eukaryotes and organelles, comparative genomics, population genetics, the microbiome, and human evolution. Prerequisites: 214, 215, or CHM 236. One three-hour seminar. Note that students new to either evolution or genetics will find 309 more appropriate. L. Landweber

EEB 321 Ecology: Species Interactions, Biodiversity and Society   Fall

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. R. Pringle

EEB 322 Advanced Ecology   Not offered this year

An advanced overview of the structure of ecological communities, particularly temperate and tropical forests. Emphasis will be on factors governing species diversity and abundance on both local and global scales. Other topics will include the impact of humans on biodiversity at global scales, and the effects of biodiversity on the regulation of climate and the cycling of key elements such as carbon and nitrogen. Prerequisite: 321; one year of calculus recommended. Two 90-minute lectures, one preceptorial. S. Pacala

EEB 323 Integrative Dynamics of Animal Behavior   Not offered this year

An exploration of the fundamental principles underlying the organization and function of animal behavior. This course will examine how complex actions emerge from simple rules. Since forces shaping behavior naturally cross scales and disciplinary boundaries, this course will draw on information from neuroscience, evolutionary biology, ecology, physiology, genetics, and the biology of complex systems. Two 90-minute lectures, one preceptorial. Staff

EEB 324 Theoretical Ecology   Spring QR

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. S. Levin

EEB 326 Human Genomics: The Past, Present and Future of the Human Genome (see ISC 326)

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

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. A. Graham

EEB 328 Ecology and Epidemiology of Parasites and Infectious Diseases   Spring STL

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. 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. Staff

EEB 336 The Diversity of Brains (see PSY 336)

EEB 338 Tropical Biology (also LAS 351)   Spring STL

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. Y. Basset

EEB 346 Biology of Coral Reefs   Spring STL

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. S. Pacala

EEB 350 Vertebrate Tropical Ecology   Not offered this year

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. Staff

EEB 355 Introduction to Statistics for Biology (also MOL 355)   Not offered this year QR

An applied introduction to probability and statistical methods in biology. Topics include data visualization, parameter estimation, frequentist hypothesis testing, Bayesian hypothesis testing, useful statistical tests, linear models, and an introduction to data-driven modeling. Analyses will use an appropriate statistical computer package. J. Storey, P. Andolfatto

EEB 400 Seminar in Global Health and Health Policy (see GHP 400)

EEB 404 Natural History of Mammals   Spring STL

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: 210 or 211. D. Rubenstein

EEB 414 Genetics of Human Populations (also MOL 414)   Not offered this year

This seminar surveys the evolutionary history of modern humans and the genetic basis of variation in our species through reading and discussion of classic and contemporary primary literature. Topics include the evolutionary origins of modern human populations, signatures of natural selection in the human genome, and approaches for discovering genetic variants that affect disease susceptibility and variation in normal traits. Significant emphasis is placed on recent advances made possible by the human genome project. Prerequisites: 309, or 320, or MOL 342, or CHM 236. One three-hour seminar. P. Andolfatto

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

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. L. Hedin

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

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. L. Hedin

EEB 419 Environmental Microbiology (see GEO 417)