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Human Evolutionary Biology

Faculty of the Department of Human Evolutionary Biology

Daniel E. Lieberman, Edwin M. Lerner II Professor of Biological Sciences, Harvard College Professor (Chair)
John C. Barry, Lecturer on Human Evolutionary Biology
Terence D. Capellini, Assistant Professor of Human Evolutionary Biology
Lara Durgavich, College Fellow in Human Evolutionary Biology
Peter T. Ellison, John Cowles Professor of Anthropology (on leave spring term)
Judith F. Chapman, Lecturer on Human Evolutionary Biology
Amy Hansen, Preceptor in the Life Sciences
Katherine J. Hinde, Assistant Professor of Human Evolutionary Biology
Carole K. Hooven, Lecturer on Human Evolutionary Biology
Susan F. Lipson, Lecturer on Human Evolutionary Biology
Zarin Pearl Machanda, College Fellow in Human Evolutionary Biology
Stephanie L. Meredith, College Fellow in Human Evolutionary Biology
David Pilbeam, Henry Ford II Professor of Human Evolution (Director of Undergraduate Studies)
Linda M. Reynard, Lecturer on Human Evolutionary Biology
Alexandra Rosati, Visiting Assistant Professor of Human Evolutionary Biology
Maryellen Ruvolo, Professor of Human Evolutionary Biology, Harvard College Professor (Director of Graduate Studies)
Tanya M. Smith, Associate Professor of Human Evolutionary Biology
Noreen Tuross, Landon T. Clay Professor of Scientific Archaeology
Anna G. Warrener, Lecturer on Human Evolutionary Biology
Richard W. Wrangham, Ruth Moore Professor of Biological Anthropology (on leave spring term)
Katherine Diane Zink, College Fellow in Human Evolutionary Biology

Other Faculty Offering Instruction in the Department of Human Evolutionary Biology

Andrew A. Biewener, Charles P. Lyman Professor of Biology (on leave fall term)
Stacey A. Combes, Associate Professor of Organismic and Evolutionary Biology
George V. Lauder, Henry Bryant Bigelow Professor of Ichthyology and Curator of Ichthyology in the Museum of Comparative Zoology

Affiliates of the Department of Human Evolutionary Biology

David E. Reich, Professor of Genetics (Medical School)

Human Evolutionary Biology provides a general foundation in human and organismic biology as part of the Life Sciences cluster of concentrations. It addresses why humans and primates are the way they are from an evolutionary perspective. Understanding the biological bases for the behavioral and physical traits that distinguish humans from other primates is one of the great challenges of modern biology, and is the focus of Human Evolutionary Biology. Students interested in addressing questions about human and non-human primate cognition from the perspective of human evolutionary biology also may pursue a special program of study affiliated with the University-wide Mind, Brain, and Behavior Initiative. For concentration requirements, see the Department of Human Evolutionary Biology website.

Primarily for Undergraduates

*Human Evolutionary Biology 91r. Supervised Reading and Research
Catalog Number: 3631
David Pilbeam
Half course (fall term; repeated spring term). Hours to be arranged. EXAM GROUP: Fall: 4
Special study of selected topics in human evolutionary biology, given on an individual basis and directly supervised by a member of the Human Evolutionary Biology Faculty.
Note: May be taken for a letter grade or Pass/Fail. Signature of faculty supervisor required.

*Human Evolutionary Biology 97. Sophomore Tutorial in Human Evolutionary Biology
Catalog Number: 2205
David Pilbeam
Half course (spring term). Hours to be arranged.
An introduction to the issues and methods of human evolutionary biology, focusing on evolutionary theory, the concept of adaptation, and their application to human evolution. Weekly readings and discussions, with biweekly writing assignments that integrate major course themes.
Note: Required of and limited to Human Evolutionary Biology concentrators.

*Human Evolutionary Biology 99a. Tutorial—Senior Year
Catalog Number: 2840
Katherine D. Zink
Half course (fall term). Hours to be arranged. EXAM GROUP: 3
Research and writing of the Senior Thesis.
Note: Limited to honors candidates. Signature of the faculty adviser required.

*Human Evolutionary Biology 99b. Tutorial - Senior Year
Catalog Number: 61326
David Pilbeam
Half course (spring term). Hours to be arranged.
Research and writing of the Senior Thesis.
Note: Limited to honors candidates. Signature of the faculty adviser required.

Cross-listed Courses

*Freshman Seminar 44s. Neanderthals and Human Evolutionary Theory
Life Sciences 1b. An Integrated Introduction to the Life Sciences: Genetics, Genomics, and Evolution
Life Sciences 2. Evolutionary Human Physiology and Anatomy
Science of Living Systems 16. Human Evolution and Human Health

For Undergraduates and Graduates

*Human Evolutionary Biology 1210. Research in Comparative Biomechanics: Seminar
Catalog Number: 11259
Andrew A. Biewener, Stacey A. Combes, George V. Lauder, Daniel E. Lieberman, and Anna G. Warrener
Half course (spring term). Hours to be arranged.
Introduces students to experimental techniques used to investigate the structure and physiology of humans and other animals. Each instructor offers research projects that are undertaken in their laboratory (limit 5 students per instructor). Students meet to introduce their project, discuss their work and progress, and to present their final results. An extensive commitment of time in the laboratory is required. Grades are based on the work completed, the oral presentation, and a short research paper.
Note: Laboratory safety session required.
Prerequisite: Life Sciences 2 or Organismic and Evolutionary Biology 102 or equivalent preferred.

[Human Evolutionary Biology 1275. Walk This Way: Sex Differences in Locomotion]
Catalog Number: 85913 Enrollment: Limited to 8.
Anna G. Warrener
Half course (fall term). Tu., Th., 10–11:30.
This course takes a broad look at how differences in male and female structure and physiology affect locomotion and movement. The first half of the course will include lectures and discussions introducing students to the biological determinants of sex, anatomical variation, and biomechanics. Students then participate in lab-based data collection and biomechanics analysis focusing on how men and women are different and similar in movement profiles. Grades are based on exams, discussion and lab participation, and a short paper and presentation.
Note: Expected to be given in 2015–16. Lab safety overview and human subjects training course required.
Prerequisite: Science of Living Systems 16 or Life Sciences 2 or approval of instructor.

Human Evolutionary Biology 1310. Hormones and Behavior
Catalog Number: 2265
Carole K. Hooven
Half course (spring term). M., W., F., at 11, and a weekly section to be arranged. EXAM GROUP: 14
An introduction to the interaction between hormones and behavior, emphasizing research in humans. General principles of endocrine physiology are presented. The course then focuses on how hormones affect the brain and body in early development and later in adulthood, and the relationship of hormones to sex and gender. We will explore human reproduction, energy metabolism, mating and sexuality, parental behavior, stress, and dominance interactions.
Note: This course is a prerequisite for Human Evolutionary Biology 1418.

[*Human Evolutionary Biology 1312. Human Sexuality: Research and Presentation Seminar]
Catalog Number: 5008 Enrollment: Limited to 15.
Judith Flynn
Half course (fall term). Th., 3–5.
An examination of human sexuality from a scientific perspective. Students will read and present primary scientific literature that highlights current research on a variety of topics including: sexual development, gender identity, sexual orientation, cross cultural variations in mating systems, promiscuity, the evolution of monogamy, sexual attraction, sexual communication, including an exploration of the existence of human pheromones, libido and sexual dysfunction.
Note: Expected to be given in 2015–16.

*Human Evolutionary Biology 1313. Stress: Research and Presentation Seminar
Catalog Number: 27108 Enrollment: Limited to 15.
Judith Flynn
Half course (spring term). Th., 3–5. EXAM GROUP: 16
An examination of stress from a scientific perspective with a focus on stress research in mammals, especially primate and humans. A writing and speaking intensive seminar that will explore the basics of the stress response, physiological effects of the stress and factors that affect stress responsiveness, such as perinatal and early life effects, social support, outlets for frustration and coping skills. The relationship between stress and disease will also be explored. Scientific studies of the effectiveness of modalities of stress reduction will also be discussed. Students will present primary scientific literature that highlights current research on a variety of topics in the field.

Human Evolutionary Biology 1329. Sex, Love and War: The Evolution of Human Behavior
Catalog Number: 89352
Richard W. Wrangham and Katherine J. Hinde
Half course (fall term). M., W., 11:30–1. EXAM GROUP: 18
This introductory course is designed to familiarize students with the behavioral ecology of humans from an evolutionary perspective, including evolutionary psychology and dual inheritance theory. We will survey behavioral diversity and consistency across human societies, and we will gain insights into the adaptive significance of human behavior and social organization by reference to social dynamics in other species. Topics to be covered include cooperation, aggression and warfare, dominance and hierarchy, mating and pair-bonds, parenting, social learning, culture and religion.
Note: This course is most relevant for freshman and sophomores who have not yet declared a concentration in Human Evolutionary Biology, or for HEB concentrators who have not taken courses in the evolution of behavior.

Human Evolutionary Biology 1330. Primate Social Behavior
Catalog Number: 4332
Zarin P. Machanda and Stephanie L. Meredith
Half course (spring term). M., W., F., at 1, and a weekly section to be arranged. EXAM GROUP: 8
A review of the behavioral interactions in natural primate populations, drawing on experimental, observational, and theoretical studies. Discussion of ecological, physiological, and developmental bases of primate social behavior, with special attention to the evolution of patterns of behavioral interactions among individuals of different age, sex, relatedness, and status. Topics include sexual conflict, sexual selection, and mating systems; care of offspring and other aid-giving; manipulative and cooperative aspects of communication; competition, dominance, and territoriality; and the evolution of social relationships.

[Human Evolutionary Biology 1335. Behavioral Ecology of Chimpanzees]
Catalog Number: 32284
Zarin P. Machanda
Half course (spring term). W., 1–4.
An advanced seminar on current topics in behavioral ecology research of chimpanzees and bonobos. Topics will include: foraging, dominance, cooperation, adolescence, reproductive strategies, culture, ranging, cognition, molecular ecology, and relationships. We will discuss behavioral flexibility of chimpanzees between different communities across Africa and learn how to collect and analyze behavioral data. We will compare the behavior of chimpanzees and bonobos with that of humans and examine how these species might serve as models for human evolution.
Note: Expected to be given in 2015–16.
Prerequisite: Human Evolutionary Biology 1330 or Science B-29 or permission of instructor.

Human Evolutionary Biology 1351. Reproductive Ecology
Catalog Number: 3408
Lara Durgavich
Half course (fall term). Tu., Th., 10–11:30. EXAM GROUP: 12
A course on the physiological ecology and evolutionary biology of human and primate reproduction. Topics covered include gamete production, gestation, birth, lactation, reproductive maturation, mature reproductive function, aging and senescence.
Prerequisite: Life Sciences 2 or Human Evolutionary Biology 1310.

Human Evolutionary Biology 1366. Mating Strategies
Catalog Number: 62507 Enrollment: Limited to 15.
Stephanie L. Meredith
Half course (spring term). W., 2–5. EXAM GROUP: 18
In this advanced seminar, we will examine the selection pressures that drive animals (including the human animal) to make particular mating decisions. We will engage in a broad, comparative exploration of the diversity of mating strategies across the animal kingdom, paying particular attention to primates, in order to ground our understanding of human mating strategies in an evolutionary perspective. Topics to be covered include the evolution of: sex, paternal care, sex-role reversal, social monogamy versus sexual monogamy, sexual coercion, homosexual behavior, and frequency dependent mating strategies.
Note: Human Evolutionary Biology concentrators will have priority, if space is limited.
Prerequisite: Human Evolutionary Biology 97 or Human Evolutionary Biology 1330 or Organismic and Evolutionary Biology 57 or permission of instructor.

[Human Evolutionary Biology 1380. Behavioral Biology of Women]
Catalog Number: 8721 Enrollment: Limited to 30.
Lara Durgavich
Half course (fall term). Tu., Th., 10–11:30.
This course is an exploration of female behavioral biology from an evolutionary and biosocial perspective. We will focus on physiological, ecological, and social aspects of women’s development from puberty, through reproductive processes such as pregnancy, birth and lactation, to menopause and aging. We will also explore female life-history strategies in a variety of cultural settings. Topics include cognitive and behavioral differences between men and women and male and female reproductive strategies. Examples are drawn primarily from traditional and modern human societies; data from studies of nonhuman primates are also considered.
Note: Expected to be given in 2015–16. Priority given to Human Evolutionary Biology concentrators.

[Human Evolutionary Biology 1411. Evolution and Adaptation of the Human Diet]
Catalog Number: 89118
Noreen Tuross and Richard W. Wrangham
Half course (fall term). M., W., F., at 11; Tu., at 2.
Within and across cultures people adopt widely varying diets, yet as a species, our foods are characteristically human. In this course we ask what is the fundamental nature of the human diet, what constrains it, how people adapt to different diets, and how the human diet evolved from those of our primate ancestors.
Note: Expected to be given in 2015–16.

[Human Evolutionary Biology 1416. The Neurobiology of Sociality: Seminar]
Catalog Number: 42215 Enrollment: Limited to 15.
Katherine J. Hinde
Half course (spring term). W., 2:30–5:30.
Recent research has illuminated the neural mechanisms underpinning sociality and social behavior in humans and other animals. In this seminar we will discuss publications that address modifications to neural structure and function as a result of behavioral specializations among taxa in relation to their social complexity or among individuals within species as a function of their social condition. This course will emphasize the value of approaching neurobiology from an evolutionary perspective and understanding the selective pressures that have shaped our mind, brain, and behavior.
Note: Expected to be given in 2015–16. Can be taken by Human Evolutionary Biology concentrators as a Junior Research Seminar. Open to graduate students.
Prerequisite: Previous or concurrent registration in Molecular and Cellular Biology 80 strongly recommended.

*Human Evolutionary Biology 1418. Endocrinology and Behavior: Research Seminar
Catalog Number: 1437 Enrollment: Limited to 8.
Susan F. Lipson
Half course (fall term). M., W., (F.), at 11, and a weekly laboratory either M. or W., 2-5. EXAM GROUP: 18
An introduction to laboratory techniques and research design in behavioral endocrinology. Students conduct pilot research projects.
Note: Preference given to Human Evolutionary Biology concentrators. Lab safety training required (after enrollment).
Prerequisite: Human Evolutionary Biology 1310 or Life Sciences 2 or with permission of instructor.

Human Evolutionary Biology 1419. Laboratory Methods in Human Evolutionary Biology
Catalog Number: 98922 Enrollment: Limited to 10.
Linda M. Reynard
Half course (spring term). W., 1–4. EXAM GROUP: 8
An introduction to laboratory methods in Human Evolutionary Biology. We will use state-of-the-art equipment and techniques to explore how humans metabolize caffeine, starch, and alcohol. Topics include quantitative pcr and genetic analysis, immunological methods, enzyme kinetics, chromatography, and measurement of isotope ratios in tissues.
Prerequisite: Life Sciences 1 and Life Sciences 1b.

Human Evolutionary Biology 1420. Human Evolutionary Anatomy
Catalog Number: 6233
Tanya M. Smith
Half course (spring term). M., W., F., at 10, and a weekly section to be arranged. EXAM GROUP: 5
How did the human body evolve, and how does it develop, grow and function? This course provides an integrative regional overview of human anatomy, with an emphasis on the musculo-skeletal system, and a comparative approach to the evolution of modern anatomy. Additional topics include: skeletal and dental development; gross anatomy of the nervous and circulatory systems; comparative limb anatomy; and comparative cranial anatomy.
Prerequisite: Life Sciences 2 or with permission of instructor.

Human Evolutionary Biology 1421. Teeth
Catalog Number: 8758 Enrollment: Limited to 6.
Tanya M. Smith
Half course (fall term). W., 3–5:30. EXAM GROUP: 6
Teeth are one of the best preserved and most commonly-recovered elements in fossil assemblages. This seminar will focus on ways in which dental remains may inform studies of primate growth and development, ecology, and health. Students will read and discuss current scientific literature, engage in histological studies in the Dental Hard Tissue Laboratory, and conduct pilot research projects.
Note: Preference given to Human Evolutionary Biology junior and senior concentrators.

Human Evolutionary Biology 1424. Human Health in Evolutionary and Anthropological Perspective
Catalog Number: 85528 Enrollment: Limited to 12.
Lara Durgavich
Half course (fall term). F., 10–1. EXAM GROUP: 5
This course applies a Darwinian perspective to explore the ultimate causes of human disease, and uses the tenets of evolutionary theory to explain variability in the health of individuals and populations. In addition, we will examine the role that environmental conditions, economic factors, and sociocultural practices play in shaping modern patterns of human health and disease. Topics will include human-pathogen coevolution, diet and nutrition, reproductive health, mental illness, and senescence.
Note: Priority given to Human Evolutionary Biology concentrators
Prerequisite: Introductory course that covers the basics of evolutionary theory.

*Human Evolutionary Biology 1434. Primate Behavior Lab
Catalog Number: 19021 Enrollment: Limited to 8.
Stephanie L. Meredith
Half course (fall term). Tu., Th., 10–11:30. EXAM GROUP: 12
In this class, we will collaboratively design a data collection protocol to answer questions of interest to both students and zookeepers regarding the Franklin Park Zoo gorillas, collect behavioral data at the Franklin Park Zoo (students will need to be able to commit to 5-6 hours of data collection during those weeks), analyze our behavioral data, write up study results in the format of a publishable scientific paper, and create a scientific meetings-style poster presentation of study results to be shared with the staff of the Franklin Park Zoo.
Note: Signature of instructor is required to enroll. Enrollment will be limited to 8students. Class meeting time to be changed as needed.
Prerequisite: Suggested prerequisites are HEB 1330, HEB 1329 or equivalent.

[Human Evolutionary Biology 1435r. Primate Ecology and Evolution]
Catalog Number: 83093
John C. Barry
Half course (fall term). Tu., Th., 10-11:30, and a weekly section to be arranged.
The lecture-seminar course will explore how primates have evolved and adapted. We will examine both living and fossil primates and discuss the degree to which "environmental" change is implicated in evolution. Goals include providing direct, hands-on experience with fossils and fostering an understanding of the strengths and limitations of the fossil record. Topics will include adaptations for food harvesting and processing, life history strategies, sexual dimorphism, and locomotion.
Note: Expected to be given in 2015–16.
Prerequisite: None, but Life Sciences 2, Science of Living Systems 16, Human Evolutionary Biology 1420, or Human Evolutionary Biology 1330 would be helpful.

*Human Evolutionary Biology 1451. Primate Functional Genetics and Genomics
Catalog Number: 99399 Enrollment: Limited to 12.
Terence D. Capellini
Half course (spring term). Hours to be arranged.
Recent advances in genetics, genomics, and developmental biology are improving our understanding of human and non-human primate biological traits. These disciplines, when incorporated into a multi-faceted context, can reveal the mechanistic basis of evolutionary adaptations. This seminar is designed to investigate and critically evaluate foundational and novel research in primates (and other organisms) that employs the tools of these trades. In doing so, students are exposed to an integrative perspective upon which to explore classic and modern questions in functional biology.
Prerequisite: Life Sciences 1b or equivalent genetics/genomics course.

*Human Evolutionary Biology 1463. Molecular Evolution of the Primates
Catalog Number: 3359 Enrollment: Limited to 12.
Maryellen Ruvolo
Half course (fall term). Th., 1–4. EXAM GROUP: 8
Introduction to the primates, emphasizing their molecular evolutionary history and the forces that mold their genomes. Topics include the neutral theory of molecular evolution, molecular clock concept and its applications, evolution of multigene families, relationships between primate morphological and molecular evolution, molecular convergences, evidence for horizontal gene transfer in primate genomes, and evolution of simian and human immunodeficiency viruses, color vision genes.
Prerequisite: Life Sciences 1b.

Human Evolutionary Biology 1480. Human Evolution through Developmental Change - (New Course)
Catalog Number: 28385
Terence D. Capellini and David Pilbeam
Half course (spring term). Tu., Th., 10–11:30. EXAM GROUP: 12
Humans and our primate relatives display marked variation in biological traits. This variation results from natural selection operating on pre- and post-natal developmental mechanisms. While these mechanisms remain mostly unknown, recent advances in genetics, genomics, and developmental biology now allow us to begin to understand how evolutionary processes influence and are influenced by underlying developmental and genetic organization. This course explores these inter-relationships in the context of the primate paleontological record. We focus on the evolution of the cranium, dentition, axial skeleton, and limbs, and present studies that cast light on the mechanisms that underlie major transitions in human evolution.
Prerequisite: Life Sciences 1b

Human Evolutionary Biology 1490r. Primate Evolution
Catalog Number: 7376
David Pilbeam and John C. Barry
Half course (fall term). Tu., Th., 10–11:30 and a weekly section, W., 1:30-3:30. EXAM GROUP: 12
A lecture/discussion course on primate evolution from a paleontological perspective. Following a survey of major primate groups as adaptive radiations, the hominoid fossil record will be reviewed within the context of the mammalian record, a particular focus being the relationship between adaptive, faunal, and climate change. Systems that can be inferred from the fossil record (for example, positional and foraging behaviors) will be discussed comparatively.
Note: No final exam; research paper required. Can be taken by Human Evolutionary Biology concentrators as a Junior Research Seminar. Introductory courses in paleoanthropology, evolution, genetics, or anatomy helpful.

Human Evolutionary Biology 1495. The Head
Catalog Number: 83815 Enrollment: Limited to 12.
Katherine D. Zink
Half course (fall term). W., 2–5. EXAM GROUP: 7
How and why does the human head look the way it does? How does the head develop, and what is the interplay between this development and how the head evolves? Why are human brains so anomalously large, while our faces are quite small? In this seminar, we will explore the evolution and natural history of the head, using the comparative anatomy of our primate and hominin relatives as a guide.
Prerequisite: Life Sciences 2 or Human Evolutionary Biology 1420 recommended, but not required.

Human Evolutionary Biology 1500. Building Babies: Developmental Trajectories from Conception to Weaning
Catalog Number: 93737 Enrollment: Limited to 8.
Katherine J. Hinde
Half course (spring term). W., 2:30–5:30. EXAM GROUP: 18
Research on human and non-human primate developmental trajectories has grown exponentially among numerous disciplines including evolutionary anthropology, psychobiology, nutrition, behavioral biology, and neuroscience. The seminar will cover the mechanisms, function, and evolution of human and non-human primate development from conception through pregnancy and lactation. Areas of development to be included will be somatic growth, immunology, behavioral/social interactions, neurobiology/cognition/learning, and metabolic processes.

[*Human Evolutionary Biology 1530. Hominid Paleontology and Evolution]
Catalog Number: 52879 Enrollment: Limited to 12.
David Pilbeam and John C. Barry
Half course (fall term). Tu., Th., 10–11:30 and a weekly section.
A lecture, discussion, and research course using cast and comparative collections, focusing on important issues in hominid paleobiology: ape ancestors, human ancestors, and early hominin radiations; earliest Homo; neandertals and modern humans; the role of environmental change. Can be taken as a research seminar.
Note: Expected to be given in 2015–16.
Prerequisite: Life Sciences 2 or Science of Living Systems 16 or Human Evolutionary Biology 1420 or with permission of instructor.

Human Evolutionary Biology 1540. Human Migration
Catalog Number: 68708 Enrollment: Limited to 10.
Noreen Tuross
Half course (spring term). Tu., 1–3. EXAM GROUP: 1
The course will explore human migration at several scales, time depths and data sources, including the movement of humans out of Africa and the complex movements of the first farmers across Europe. We will explore the impacts that climates and disease burden have had on human migrations, and discuss recent movements of people and the reasons for migratory behavior in humans. in addition, a personal migration story will be developed by the class.

Human Evolutionary Biology 1590. Ancient Biomolecules - (New Course)
Catalog Number: 93588 Enrollment: Limited to 30.
Noreen Tuross and Linda M. Reynard
Half course (fall term). Tu., Th., 1:30-3, and a weekly section to be arranged. EXAM GROUP: 8
How do we know where and how people migrated? How old is this ruin? Did ancient humans eat a lot of meat? This course will explore the known, the unknown, and the unknowable in the study of ancient biomolecules and critically evaluate the current literature and the accompanying press reports. The course couples topics about the past in which ancient biomolecules are used with an examination of the methods employed. The format of the course will be lecture and case study.
Note: This course, when taken for a letter grade, meets the General Education requirement for Empirical and Mathematical Reasoning.
Prerequisite: There are no prerequisites, but you can expect to do more work if you have not taken Life and Physical Sciences A or Life Sciences 1a or had a strong high-school biology course.

Cross-listed Courses

*OEB 121a. Research in Comparative Biomechanics: Seminar

Primarily for Graduates

*Human Evolutionary Biology 2312. Current Topics in Human Evolutionary Genetics
Catalog Number: 1175
Maryellen Ruvolo
Half course (spring term). Hours to be arranged.
Critical reading of current literature on the genetics of living humans and discussion of evolutionary implications.
Note: Open to first and second year graduate students in Human Evolutionary Biology.

[Human Evolutionary Biology 2335. Introduction to Mathematical Modeling in Human Evolutionary Biology]
Catalog Number: 73853 Enrollment: Open to undergraduates with instructor’s permission.
Peter T. Ellison
Half course (spring term). Th., 2–5.
This course will introduce students to the use and interpretation of the major forms of mathematical models used in contemporary evolutionary biology, including dynamical population models, game theoretic models, and agent-based models. Sophisticated mathematical training and ability are not required, but some familiarity with first year calculus is assumed. The course will include an introduction to relevant software packages for mathematical analysis and simulation. Application of mathematical models to contemporary topics and debates such as the evolution of cooperation and life history evolution will be explored. Students will develop a model of their own as a class project.
Note: Expected to be given in 2015–16.
Prerequisite: Mathematics 1a,1b or equivalent strongly recommended.

Human Evolutionary Biology 2430. Behavioral Biology Seminar
Catalog Number: 3777
Katherine J. Hinde
Half course (fall term). Hours to be arranged. EXAM GROUP: 9
Reading and discussion of current research in the behavioral ecology of humans and nonhuman primates. Emphasis placed on comparative and interdisciplinary approaches.
Note: Required of entering graduate students in Human Evolutionary Biology. Open to other graduate students. Limited to graduate students.

Human Evolutionary Biology 2460. Issues in Human Evolution
Catalog Number: 1073
David Pilbeam
Half course (spring term). W., 1–2:30. EXAM GROUP: 8
A discussion course for graduate students in Human Evolutionary Biology. Topics will include origins of hominids, radiation of hominins, origins of the genus Homo, and origins of Homo sapiens.
Note: May be taken while auditing Science of Living Systems 16.

*Human Evolutionary Biology 2590. Recent Topics in Ancient Biomolecules - (New Course)
Catalog Number: 34717
Noreen Tuross
Half course (fall term). Hours to be arranged. EXAM GROUP: 17
Graduate seminar to accompany undergraduate lecture course on ancient biomolecules.
Note: Primarily for graduate students, but undergraduates welcome with instructor’s permission.

Cross-listed Course

Graduate Courses of Reading and Research

*Human Evolutionary Biology 3000. Reading and Research
Catalog Number: 99167
John C. Barry 1892, Terence D. Capellini 7274, Peter T. Ellison 7413 (on leave spring term), Katherine J. Hinde 6956, Daniel E. Lieberman 3980, Susan F. Lipson 1969, David Pilbeam 7224, Maryellen Ruvolo 2512, Tanya M. Smith 6147, Noreen Tuross 4845, and Richard W. Wrangham 2349 (on leave spring term)
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 1; Spring: 8
Special reading in selected topics under the direction of members of the department.
Note: Consult the appropriate member of the department.

*Human Evolutionary Biology 3001. Reading for General Examination
Catalog Number: 47645
Members of the Department
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 7; Spring: 18
Individual reading in preparation for the general examination for the doctoral degree.
Note: Restricted to candidates for the doctoral degree and ordinarily to those who have completed at least one year in residence.

*Human Evolutionary Biology 3200. Graduate Seminar in Human Evolutionary Biology
Catalog Number: 31571
David Pilbeam 7224 and Terence D. Capellini 7274
Half course (fall term). W., 10–11:30. EXAM GROUP: 1
Proseminar for Human Evolutionary Biology graduate students. Discussion of adaptations and the process of adaptation using examples from various areas of human evolutionary biology.
Note: Open to graduate students in Human Evolutionary Biology.

*Human Evolutionary Biology 3310. Experimental Methods
Catalog Number: 9602
Members of the Department
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 1; Spring: 8

*Human Evolutionary Biology 3320. Advanced Laboratory and Dissertations
Catalog Number: 62752
Members of the Department
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 6; Spring: 17

*Human Evolutionary Biology 3337. Advanced Laboratory Methods in Human Endocrinology
Catalog Number: 5345
Susan F. Lipson 1969
Half course (fall term; repeated spring term). . EXAM GROUP: 15
Note: Intended for graduate students engaged in laboratory research on human endocrinology.

*Human Evolutionary Biology 3350. Laboratory Methods in Primate and Human Nutrition
Catalog Number: 62293
Richard W. Wrangham 2349 (on leave spring term)
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 9; Spring: 2
Independent laboratory study in the biochemical analysis of plant and animal foods, and of human and animal digestive physiology and feeding behavior.

*Human Evolutionary Biology 3400. Advanced Reading and Research
Catalog Number: 77859
Members of the Department
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 6; Spring: 17
Note: Consult the appropriate member of the department.

*Human Evolutionary Biology 3500. Direction of the Doctoral Dissertation
Catalog Number: 26337
Members of the Department
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 17; Spring: 13
Note: Consult the appropriate member of the department.

*Human Evolutionary Biology 3595. Laboratory Methods in Evolutionary Genetics
Catalog Number: 7934
Maryellen Ruvolo 2512
Half course (fall term; repeated spring term). . EXAM GROUP: Fall: 9; Spring: 2
Note: Limited to graduate students conducting doctoral dissertation research.

*Human Evolutionary Biology 3600. Current Issues in Human Evolutionary Biology
Catalog Number: 9373
Members of the Department
Half course (fall term; repeated spring term). Tu., at 12. EXAM GROUP: Fall: 11; Spring: 7
Weekly seminars in human evolutionary biology.