Research in the Department of Human Evolutionary Biology asks the question "Why are humans the way we are?" using evolutionary and comparative approaches to human biology and behavior. The faculty view several major, interrelated research questions as most important to the study of human evolutionary biology. First, what selective forces operated at different times in primate and human evolution, including today, to make humans the way we are behaviorally, psychologically, anatomically, physiologically, and genetically? Second, what are the genetic versus environmental underpinnings of the human phenotype- both behavioral and physical-- in terms of fixed differences from other primates and of polymorphic differences that vary among humans? Third, how do the evolutionary bases of important aspects of human biology interact with social and cultural factors in shaping problems we currently face such as violence, poverty, overpopulation, and disease? In order to address these questions, the department integrates theoretical and metodological tools from other sciences as needed; these currently include behavioral ecology of human and non-human primates, genetics and genomics, developmental and functional anatomy, paleontology, physiology, systems biology, psychology, economics and anthropology. We also incorporate laboratory and field-based research.

The objective of the PhD program is to provide students with comprehensive training taking a strongly evolutionary and comparative approach to human adaptations and their evolution. Faculty research spans a broad range of approaches aimed at understanding the evolution of humans and their closest primate relatives. Our interdisciplinary approach includes field and/or laboratory programs in endocrinology, human behavioral biology and ecology, ape behavioral ecology and biology, human and primate paleobiology, experimental biomechanic and developmental anatomy, experimental physiology, microbiology and metagenomics, cultural evolution, primate comparative cognition, and the genetics and genomics of humans and primates.

We are open to interdepartmental Ph.D. programs. We have strong collaborative links to faculty in the departments of Anthropology, Organismic and Evolutionary Biology, Psychology, Economics, and Harvard Medical School. Primary advisors of graduate students must be faculty members within the Department of Human Evolutionary Biology.

The department is located mainly on the fifth floor of the Peabody Museum, Harvard University, 11 Divinity Avenue, Cambridge, Massachusetts 02138 and houses laboratory research facilities in genetics, paleontology, skeletal and dental biology, experimental biomechanics, endocrinology, nutritional analysis, microbiology, and isotopic and materials analysis. Other Harvard resources include: FAS Center for Systems Biology, Museum of Comparative Zoology (including the Concord Field Station in Concord, Massachusetts), the Center for Nanoscale Systems, the Peabody Museum’s extensive human and non-human skeletal collections, and extensive facilities for human and non-human experimental research.

Degrees Offered

The graduate program in human evolutionary biology is a PhD degree program. There is no terminal masters degree program in human evolutionary biology. Students apply to and are admitted to the doctoral degree program only.

Since the program’s principal objective is to prepare students for college or university teaching or research, for which the doctoral degree is required, the master’s degree in human evolutionary biology is usually not taken as an end in itself. While a student enrolled in the doctoral program may earn a masters degree en route to their doctoral degree, as the student qualifies for it, the masters degree is not normally taken as an end in itself, unless an enrolled student is unable or chooses not to continue to the dissertation phase of the doctoral program.

Admission Requirements

Our PhD program is small, admitting each year only a few students with good backgrounds in evolutionary biology and with research interests that correspond with those of our faculty, which would include faculty who hold the rank of Assistant Professor, Associate Professor, or Professor. It would not include faculty who hold the rank of College Fellow, Lecturer, or Preceptor. Applicants must hold a Bachelor’s degree, ordinarily with distinction, and take the verbal, quantitative, and analytical aptitude tests of the Graduate Record Examination (GRE) General Test. Successful applicants normally have strong backgrounds in math and sciences, particularly in biology, some acquaintance with statistics, and some laboratory or field research experience. We look for students with high grade-point averages, strong letters of recommendations from professors, and high GRE scores. Application for admission is submitted at

The department considers a period of five or six years in residence to be the norm for PhD candidates. For financial residence requirements, See the GSAS Admissions website.

Financial Aid

Students admitted to the PhD program receive full tuition and ten months of living support for the first four years and a final year of dissertation completion support. In the first two years they receive their living support as stipends; in the third and fourth years, as teaching or research fellowships. They also receive two-month summer research awards for the summers following the first four years in graduate school. Dissertation completion support is available as soon as the student is prepared to finish the dissertation, ordinarily in the fifth or sixth year. Progress is reviewed annually, and financial awards are contingent upon students making satisfactory progress in the program.

Prospective graduate students are urged to apply for outside fellowships that offer tuition and stipend support during graduate school. These include the National Science Foundation Graduate Research Fellowship, the Ford Foundation Diversity Fellowship, and the U.S. Department of Education’s Jacob K. Javits Fellowship. Application deadlines for these fellowships are in the fall, well before Harvard’s admissions deadline. Eligible applicants are encouraged to investigate these funding opportunities early in the application season.

Students in the department are also eligible for summer or term-time research awards and traveling fellowships funded by Harvard University. They also receive research support from the National Science Foundation, Wenner-Gren Foundation for Anthropological Research, or L.S.B. Leakey Foundation.

Degree Requirements


Two years minimum of full-time study (up to 16 half-courses or the equivalent). The department considers a period of five or six years in residence to be the norm for PhD candidates.

Appointment of Advisors

Upon admission, students are assigned two first-year faculty advisors based upon compatibility of research interests. In the week before fall term begins, first-year students will meet with their assigned faculty advisors to plan a program of study that takes into account their previous training and current academic interests. Each student’s program of study must receive the approval of his or her advisors.

First Two Years

During the first two years, students complete course requirements and begin to outline their PhD plans. Normally, students will take at least eight half-courses (semester courses) in human evolutionary biology and related areas during their first two years in residence. These will include a proseminar in evolutionary theory and a minimum of three courses out of five identified primary areas: human evolution, genetics, human anatomy and biology, behavior and ecology of humans, behavior and ecology of primates. Successful completion of the proseminar and the three area courses within the first two years constitutes a major portion of the qualifying examination process.

In addition, students must submit a draft of a research proposal (the "mock NSF") and be examined orally by the faculty. The oral examination is based on the research proposal, which students develop in consultation with their advisors, as well as their command of relevant areas of human evolutionary biology. The proposal is written in the form of a Doctoral Dissertation Improvement Grant proposal to the National Science Foundation and is circulated to the faculty at least a week before the oral examination. Students are expected to fulfill this requirement by the end of their second year in residence. Successful completion of the “mock NSF” requirement is the final step in the qualifying examination process.

Competence in appropriate statistical analysis is required by the time of the qualifying examination.

Students must acquire both theoretical grounding and technical skills. This means gaining experience with designing research projects, collecting data in the laboratory or field, and analyzing those data. To achieve this, appropriate field and/or laboratory training is required, as determined in consultation with faculty advisors. Depending upon the nature of the research to be undertaken for the PhD, the faculty may prescribe further skills, such as fluency in a field language, advanced laboratory skills, or further quantitative skills.

Years Three Through Five

After completion of the qualifying examination, the student, in consultation with his or her advisors, will select a dissertation topic. The faculty will appoint a dissertation prospectus committee, ordinarily consisting of at least three members, at least two of whom will be department members. The student, in consultation with his or her committee, will further develop the scope of the dissertation topic.

The student will submit to this committee dissertation (or thesis) prospectus that embodies the general planning of the dissertation research work and shows what contribution it will make to the field. The prospectus should give a concise statement of the problems being studied or hypotheses tested and a description of the manner in which the field or laboratory investigation will be carried out. The prospectus should conform to the format and length of an NSF Doctoral Dissertation Improvement Grant application. Ideally, it will also be a grant application. The candidate will meet with the dissertation prospectus committee to discuss the prospectus and consider any necessary revisions, including the possibility that an alternate prospectus would be required. Approval of a dissertation prospectus, including any revisions, is expected by the end of the sixth term in residence.

An approved dissertation is normally expected by the end of the twelfth term. The dissertation committee is composed of at least three readers, two of whom must be members of the Faculty of Arts and Sciences. At least one reader will be a member of the Human Evolutionary Biology faculty, and at least one reader will normally be outside that faculty. The form of the dissertation may vary depending on the student’s research, but the content should ordinarily be substantive enough to represent or to produce at least three published articles.

The dissertation defense consists of an oral presentation for a general audience followed by an oral examination attended by the dissertation committee and other interested faculty. Only after successful completion of this examination and the incorporation of any revisions required by the dissertation committee will a candidate’s dissertation be approved for submission to the Registrar.

Recent Dissertation Titles

  • Alicia Breakey, “The Life History Significance of Human Breast Milk: Immune and endocrine factors as indicators of maternal condition and predictors of infant health and growth”
  • Elizabeth Brown, “Metabolic Adaptations in Modern Human Populations: Evidence, Theory, and Investigation”
  • Rachel Carmody, "Energetic consequences of thermal and non-thermal food processing"
  • Carolyn Eng, "An Anatomical and Biomechanical Study of the Human Iliotibial Band’s Role in Elastic Energy Storage"
  • Alexander Georgiev, "Energetic costs of reproductive effort in male chimpanzees"
  • Luke Glowacki, “Incentives for war in small-scale societies”
  • Zarin Machanda, "The Ecology of Male-Female Social Relationships among East African Chimpanzees"
  • Katherine McAuliffe, "The Evolution and Development of Inequity Aversion"
  • Meredith Reiches, "Female Adolescent Energy Expenditure in The Gambia"
  • Victoria Wobber, "Comparative Cognitive Development and Endocrinology in Pan and Homo"
  • Katherine Zink, "Mechanical and thermal food processing effects on mastication and cranio-facial morphology"
  • Qu Zhang, "Understanding Genome Evolution by Comparative Genomics"

Faculty List