Integrative Biology

University of California, Berkeley

About the Program

Biological phenomena occur at various levels of structural organization, ranging from molecules to organisms, and from populations to the global ecosystem. Integrative Biology takes a whole-organism approach, extending from the genome and proteome through organismal traits (phenotypes), to communities and ecosystems. Through the coordinated study of multiple levels of biological organization over a broad range of spatial and temporal scales, Integrative Biology offers a unique approach to understanding fundamental questions concerning the evolution and maintenance of biological diversity, including organismal form and function, and ecological and ecosystem processes. This multidimensional approach underpins our graduate program, where students combine observational, experimental, and comparative approaches with the development of theory; and apply concepts and techniques from the biological sciences and other disciplines.

Integrative Biology admits students to the PhD program only.

Visit Department Website


Admission to the University

Minimum Requirements for Admission

The following minimum requirements apply to all graduate programs and will be verified by the Graduate Division:

  1. A bachelor’s degree or recognized equivalent from an accredited institution;
  2. A grade point average of B or better (3.0);
  3. If the applicant comes from a country or political entity (e.g., Quebec) where English is not the official language, adequate proficiency in English to do graduate work, as evidenced by a TOEFL score of at least 90 on the iBT test, 570 on the paper-and-pencil test, or an IELTS Band score of at least 7 (note that individual programs may set higher levels for any of these); and
  4. Sufficient undergraduate training to do graduate work in the given field.

Applicants Who Already Hold a Graduate Degree

The Graduate Council views academic degrees not as vocational training certificates, but as evidence of broad training in research methods, independent study, and articulation of learning. Therefore, applicants who already have academic graduate degrees should be able to pursue new subject matter at an advanced level without need to enroll in a related or similar graduate program.

Programs may consider students for an additional academic master’s or professional master’s degree only if the additional degree is in a distinctly different field.

Applicants admitted to a doctoral program that requires a master’s degree to be earned at Berkeley as a prerequisite (even though the applicant already has a master’s degree from another institution in the same or a closely allied field of study) will be permitted to undertake the second master’s degree, despite the overlap in field.

The Graduate Division will admit students for a second doctoral degree only if they meet the following guidelines:

  1. Applicants with doctoral degrees may be admitted for an additional doctoral degree only if that degree program is in a general area of knowledge distinctly different from the field in which they earned their original degree. For example, a physics PhD could be admitted to a doctoral degree program in music or history; however, a student with a doctoral degree in mathematics would not be permitted to add a PhD in statistics.
  2. Applicants who hold the PhD degree may be admitted to a professional doctorate or professional master’s degree program if there is no duplication of training involved.

Applicants may apply only to one single degree program or one concurrent degree program per admission cycle.

Required Documents for Applications

  1. Transcripts: Applicants may upload unofficial transcripts with your application for the departmental initial review. If the applicant is admitted, then official transcripts of all college-level work will be required. Official transcripts must be in sealed envelopes as issued by the school(s) attended. If you have attended Berkeley, upload your unofficial transcript with your application for the departmental initial review. If you are admitted, an official transcript with evidence of degree conferral will not be required.
  2. Letters of recommendation: Applicants may request online letters of recommendation through the online application system. Hard copies of recommendation letters must be sent directly to the program, not the Graduate Division.
  3. Evidence of English language proficiency: All applicants from countries or political entities in which the official language is not English are required to submit official evidence of English language proficiency. This applies to applicants from Bangladesh, Burma, Nepal, India, Pakistan, Latin America, the Middle East, the People’s Republic of China, Taiwan, Japan, Korea, Southeast Asia, most European countries, and Quebec (Canada). However, applicants who, at the time of application, have already completed at least one year of full-time academic course work with grades of B or better at a US university may submit an official transcript from the US university to fulfill this requirement. The following courses will not fulfill this requirement:
    • courses in English as a Second Language,
    • courses conducted in a language other than English,
    • courses that will be completed after the application is submitted, and
    • courses of a non-academic nature.

If applicants have previously been denied admission to Berkeley on the basis of their English language proficiency, they must submit new test scores that meet the current minimum from one of the standardized tests.

Where to Apply

Visit the Berkeley Graduate Division application page

Admission to the Program

The online Graduate Application for Admission, Fellowship and Financial Aid will be available in early September on the Graduate Division's website and will include the current deadline to apply to the program. The completed application must be submitted online and fee paid by the deadline. Be sure to allow sufficient time for your letters of recommendation and test scores to arrive by the deadline. The department reviews application for admission to our graduate program once a year. We accept applications for fall only.

Admissions Criteria

Initiating contact with faculty members; coursework; GRE general scores; letters of recommendation; degree of preparedness for graduate school; and your statement of purpose are all important factors in our review of your application.

Contact IB Faculty

It is required that you list on your application at least one faculty member in our department whose research is of interest to you. It is highly recommended that you contact them to discuss your interest in working with them. This contact is the first step in broadly defining areas of potential research focus and should be elaborated on in your statement of purpose.

Bachelor's Degree

Students admitted to the program typically have a bachelor's degree in one of the life sciences or physical sciences. However, promising students with other academic backgrounds are encouraged to apply if they have a undergraduate grounding in biology.

Grade Point Average (GPA)

Upper division or graduate GPA of 3.4 or higher is preferred. A minimum GPA of 3.0 (courses taken after the first two years) is required by the Graduate Division.

Graduate Record Examination (GRE) and TOEFL

All applicants must take the GRE general test; GRE subject test in biology or subject tests in other relevant disciplines highly recommended, but not required. No minimum GRE scores required for consideration. We will accept GRE scores taken within the last ten years. Older scores will be considered on a case by case basis. GRE Institution Code: 4833; Department Code: 0203.

For international students from countries in which the official language is not English, results of the TOEFL (Test of English as Foreign Language) are required. TOEFL exams taken before June 1, 2011 will not be accepted even if your score was reported to Berkeley.

Letters of Recommendation

Three letters of recommendation from faculty or other persons who have known you in an academic or research capacity. 

Statement of Purpose

Describe your aptitude and motivation for graduate study in your area of specialization, including your preparation for this field of study, your academic plans or research interests in your chosen area of study, and your future career goals. Please be specific about why UC Berkeley would be a good intellectual fit for you.

The statement should reflect serious intent, focus, maturity, motivation, and the ability to organize and articulate your thoughts on complex subjects.

There is no page limit restrictions although statements are typically one to two pages in length.

Personal History Statement

Please note that the personal history statement should not duplicate the statement of purpose.

Describe how your personal background informs your decision to pursue a graduate degree. Please include information on how you have overcome barriers to access opportunities in higher education, evidence of how you have come to understand the barriers faced by others, evidence of your academic service to advance equitable access to higher education for women, racial minorities, and individuals from other groups that have been historically underrepresented in higher education, evidence of your research focusing on underserved populations or related issues of inequality, or evidence of your leadership among such groups.

Research Experience

Research experience is preferred. It helps to define interest and focus, and proven success with research is a positive indicator for success in the program.

Doctoral Degree Requirements

Normative Time Requirements

  • An advanced course in evolutionary biology is the only specific course required of all graduate students. It must be taken for a letter grade during the graduate program if it was not completed during the student's undergraduate education. A student's supervisory committee may suggest courses as well.
  • Four semesters of residency as required by the Graduate Division. This means you must be registered for a minimum of four semesters. There are no departmental unit requirements for the PhD program.
  • Students are required to be a graduate student instructor (GSI) for at least two semesters and must complete INTEGBI 375.
  • A student in the PhD program must take a three hour oral qualifying examination (QE) on fields specified by their QE committee.
  • PhD candidates are required to write a dissertation based on original and independent research carried out by the student.
  • You are encouraged (but not required) to enroll in seminars in your field of specialization and present topics. Effective participation in seminars is a useful introduction to your field of specialization and may give you valuable direction for advanced study, particularly if you have not begun research activities.​


Courses Required
One advanced course in evolutionary biology (INTEGBI 160 or committee-approved alternative)
INTEGBI 375Teaching Colloquium: Graduate Student Instructor Training2
INTEGBI Electives in specialized study list - seminars and student presentations strongly advised


Integrative Biology

INTEGBI C200 Principles of Phylogenetics 4 Units

Terms offered: Spring 2018, Spring 2017, Spring 2016
The core theory and methodology for comparative biology, beginning
with issues in building phylogenetic trees, with emphases on both
morphology and molecules, and both living and fossil organisms. Also
covers the many applications of phylogenetic trees to systematics,
biogeography, speciation, conservation, population genetics, ecology,
behavior, development, functional morphology, and macroevolution
that have
revolutionized those fields. Labs are closely integrated with
lectures and cover the major algorithms and computer software used
to implement these approaches. Requirements include participation in
discussions, two exams, and a term project.

Principles of Phylogenetics: Read More [+]

INTEGBI C204 Research Reviews in Animal Behavior: Behavior Review 1 Unit

Terms offered: Fall 2018, Spring 2018, Fall 2017
This course will provide a rigorous, critical review of current research in animal behavior. Emphases will include hypothesis testing and experimental design, as well as methods of data collection and analysis. Each week, a student in the course will present original research in the form of a seminar presentation, grant proposal, or manuscript. Through discussion with seminar participants, presenters will gain critical feedback regarding their
Research Reviews in Animal Behavior: Behavior Review: Read More [+]

INTEGBI C205 Quantitative Methods for Ecological and Environmental Modeling 3 Units

Terms offered: Fall 2015, Fall 2013, Fall 2012, Fall 2011, Fall 2009
This course will review the background mathematical and statistical tools necessary for students interested in pursuing ecological and environmental modeling. Topics include linear algebra; difference equation, ordinary differential equation, and partial differential equation models; stochastic processes; parameter estimation; and a number of statistical techniques. This course will be recommended as a prerequisite for advanced
modeling courses in Integrative Biology, Energy and Resources Group, and Environmental Science, Policy, and Management.
Quantitative Methods for Ecological and Environmental Modeling: Read More [+]

INTEGBI 206 Statistical Phylogenetics 3 Units

Terms offered: Fall 2018, Fall 2012, Fall 2011
This course is aimed at students who wish to understand the evolutionary models and methods for estimating phylogenies (which are trees representing how organisms are related to one another). Topics include continuous-time Markov chains as applied in phylogenetics; maximum likelihood estimation; Bayesian estimation; the combinatorics of evolutionary trees; Markov chain Monte Carlo; distance and parsimony methods for estimating trees; optimization
strategies for finding best trees. Students will learn to write computer programs that implement many of the methods discussed in class, and apply their knowledge in a research project.
Statistical Phylogenetics: Read More [+]

INTEGBI C215 Communicating Ocean Science 4 Units

Terms offered: Spring 2018, Spring 2017, Spring 2016, Spring 2015, Fall 2014, Spring 2014, Spring 2013
For graduate students interested in improving their ability to communicate their scientific knowledge by teaching ocean science in elementary schools or science centers/aquariums. The course will combine instruction in inquiry-based teaching methods and learning pedagogy with six weeks of supervised teaching experience in a local school classroom or the Lawrence Hall of Science with a partner.
Thus, students will practice communicating scientific knowledge and receive mentoring on how to improve their presentations.
Communicating Ocean Science: Read More [+]

INTEGBI C216 Freshwater Ecology 3 Units

Terms offered: Spring 2015, Spring 2014, Spring 2013
This graduate course will combine formal lectures and discussion, with the overall goal of exposing students to general concepts in freshwater ecology. We will discuss a broad range of topics including freshwater environments and biota, natural selection and adaptive evolution, food webs and trophic cascades, cross-ecosystem linkages, and social-ecological resilience of freshwater ecosystems under global change. Upper division undergraduates
are welcome, with permission of the instructors.
Freshwater Ecology: Read More [+]

INTEGBI C217 Biomimetic Engineering -- Engineering from Biology 3 Units

Terms offered: Fall 2017, Spring 2014, Fall 2010
Study of nature's solutions to specific problems with the aim of determining appropriate engineering analogs. Morphology, scaling, and design in organisms applied to engineering structures. Mechanical principles in nature and their application to engineering devices. Mechanical behavior of biological materials as governed by underlying microstructure, with the potential for synthesis into engineered materials. Trade-offs between redundancy and
efficiency. Students will work in teams on projects where they will take examples of designs, concepts, and models from biology and determine their potential in specific engineering applications.
Biomimetic Engineering -- Engineering from Biology: Read More [+]

INTEGBI 222 Seminar in Physiological Energetics and Biomechanics 2 Units

Terms offered: Spring 2018, Spring 2015, Spring 2009
Discussion and critique of scientific literature and current topics in physiological energetics and biomechanics. Emphasis is on metabolic energetics. Topics include efficiency, energy-saving mechanisms, muscle function, oxidative stress, development in metabolic physiology and biochemistry and comparative aspects.

Seminar in Physiological Energetics and Biomechanics: Read More [+]

INTEGBI 223 Seminar on Bioenergetics and Metabolism 2 Units

Terms offered: Fall 2018, Fall 2017, Fall 2016
Immediate and long-range adaptations of the body to exercise. Physiological limits and work capacities in relation to age, sex, diet, environmental factors, and nature of activity.

Seminar on Bioenergetics and Metabolism: Read More [+]

INTEGBI C226 Isotopics 2 Units

Terms offered: Fall 2016, Fall 2015, Fall 2014
This seminar will explore current topics that employ the use of stable isotopes. Discussion topics include the areas of biology, paleontology, biogeochemistry, soil science, and atmospheric science. Students will be required to lead at least one discussion of relevant literature in the topic area.

Isotopics: Read More [+]

INTEGBI C227 Stable Isotope Ecology 5 Units

Terms offered: Spring 2016, Spring 2014, Spring 2012
Course focuses on principles and applications of stable isotope chemistry as applied to the broad science of ecology. Lecture topics include principles of isotope behavior and chemistry, and isotope measurements in the context of terrestrial, aquatic, and marine ecological processes and problems. Students participate in a set of laboratory exercises involving preparation of samples of choice for isotopic analyses, the use of the mass spectrometer
and optical analysis systems, and the anlaysis of data.
Stable Isotope Ecology: Read More [+]

INTEGBI 230 Marine Science Review 1 Unit

Terms offered: Fall 2018, Spring 2018, Spring 2017
Reports and discussion of original research in marine science.

Marine Science Review: Read More [+]

INTEGBI 232 Seminar in Biomechanics 2 Units

Terms offered: Fall 2018, Fall 2017, Spring 2017
Presentation, discussion, and critique of current literature in scientific research and current topics in comparative biomechanics which include solid and fluid mechanics, locomotion, and energetics.

Seminar in Biomechanics: Read More [+]

INTEGBI 234 Seminar on Biology of Amphibians and Reptiles 1 Unit

Terms offered: Fall 2018, Spring 2018, Fall 2017
Review of current research activity and literature concerning the biology of amphibians and reptiles.

Seminar on Biology of Amphibians and Reptiles: Read More [+]

INTEGBI 241 Advanced Topics in Endocrine-Regulated Development 3 Units

Terms offered: Spring 2018, Spring 2015, Spring 2014
This course will examine intentional endocrine disruption, such as the use of pharmaceuticals to regulate hormones in humans, livestock, and wildlife. We will also evaluate endocrine disrupting pollutants and their impacts on wildlife and humans, including their potential role in cancer.

Advanced Topics in Endocrine-Regulated Development: Read More [+]

INTEGBI 246 Seminars in Systems Biology 2 Units

Terms offered: Spring 2015, Spring 2014, Spring 2013
This course discusses seminal papers in the field of systems biology with particular emphasis on gene regulation and cell biology. The course covers the critical analysis of primary research data, computational modeling, and important theoretical concepts in systems biology. Topics vary from year to year.

Seminars in Systems Biology: Read More [+]

INTEGBI 248 Comparative Physiology and Endocrinology Seminar 1 Unit

Terms offered: Spring 2018, Spring 2017, Spring 2016
Reviews and reports of current research in vertebrate endocrinology and physiology.

Comparative Physiology and Endocrinology Seminar: Read More [+]

INTEGBI 249 Seminar on Evolutionary Genetics 1 Unit

Terms offered: Spring 2009, Spring 2008, Fall 2002
Recent developments in evolutionary genetics will be discussed in a seminar format.

Seminar on Evolutionary Genetics: Read More [+]

INTEGBI 250 Seminar in Ecology 2 Units

Terms offered: Spring 2018, Spring 2017, Fall 2014
Readings and discussion of current topics.

Seminar in Ecology: Read More [+]

INTEGBI 251 Ecological Research Reviews 1 Unit

Terms offered: Fall 2018, Spring 2018, Fall 2017
Reports and discussions of original research.

Ecological Research Reviews: Read More [+]

INTEGBI 257 Current Topics in Behavioral Physiology 2 Units

Terms offered: Spring 2010, Spring 2009, Fall 1999
Topics to vary. Report and discussion of current literature.

Current Topics in Behavioral Physiology: Read More [+]

INTEGBI 259 Advanced Paleoecology 2 Units

Terms offered: Fall 2013, Spring 2011, Spring 2009
Topics vary from year to year but will include paleoecology of major groups of organisms or major environments from population, community evolutionary, or taxonomic persepectives.

Advanced Paleoecology: Read More [+]

INTEGBI 262 Seminar in Computational Biology 1 Unit

Terms offered: Spring 2009, Fall 2008
Students will discuss original papers in the general area of computational biology and will discuss new research presented by instructors in the course and by invited speakers from other departments at UC Berkeley and from other universities and research groups.

Seminar in Computational Biology: Read More [+]

INTEGBI 263 Genetics and the Evolution of the Skeleton 2 Units

Terms offered: Spring 2016, Spring 2015, Spring 2012
In this seminar, we will explore the genetic underpinnings of vertebrate skeletal variation and review how such information is being incorporated into evolutionary and paleontological studies. Topics include quantitative genetic analyses of cranial variation and developmental genetics of the limb and dentition. This course will be tailored each semester to cover new research; therefore, students may enroll in this course multiple semeste
Genetics and the Evolution of the Skeleton: Read More [+]

INTEGBI 264 Seminar in Evolutionary Biology of the Vertebrates 1 Unit

Terms offered: Fall 2018, Spring 2018, Fall 2017
Presentation of results of original research by students, faculty, and visitors.

Seminar in Evolutionary Biology of the Vertebrates: Read More [+]

INTEGBI 265 Advanced Studies in Hominid Paleobiology 2 Units

Terms offered: Spring 2018, Spring 2017, Spring 2016
This is a graduate level course that focuses on special topics within hominid evolutionary studies. The topic for each semester will be decided upon during the first class meeting. Previous advanced training in biology, human evolutionary studies, and evolutionary theory is required.

Advanced Studies in Hominid Paleobiology: Read More [+]

INTEGBI 268 Seminar in Evolution above the Species Level 2 Units

Terms offered: Fall 2009, Fall 2006, Fall 2004
Current issues in macroevolution and paleobiology, using both neontological and paleontological data.

Seminar in Evolution above the Species Level: Read More [+]

INTEGBI 281 Seminar in Evolution 2 Units

Terms offered: Spring 2018, Spring 2017, Spring 2016
Advanced study and current literature in various fields of evolution. Topics vary from year to year.

Seminar in Evolution: Read More [+]

INTEGBI 283 Seminar in Vertebrate Evolution and Paleontology 1 Unit

Terms offered: Fall 2017, Fall 2016, Spring 2016
Presentations and discussions of original research and new literature in vertebrate evolution and paleontology. Syllabus and reading list will vary as topics change from semester to semester. Open to Undergraduate students with permission. Enrollment limit: 20.

Seminar in Vertebrate Evolution and Paleontology: Read More [+]

INTEGBI 286 Seminars in Paleontology 2 Units

Terms offered: Spring 2018, Fall 2017, Spring 2017
Advanced study and current literature in various fields of paleontology. Topics vary from year to year.

Seminars in Paleontology: Read More [+]

INTEGBI 290 Research Seminar 1 - 2 Units

Terms offered: Spring 2018, Fall 2017, Spring 2017
Advanced study in various fields of Integrative Biology. Topics will be announced in advance of each semester. Enrollment in more than one section permitted.

Research Seminar: Read More [+]

INTEGBI 291 Research Seminar 1 Unit

Terms offered: Fall 2018, Fall 2017, Fall 2016
Review and discussion of topics of current interest. Topics to vary.

Research Seminar: Read More [+]

INTEGBI 292 Integrative Biology Colloquium 0.0 Units

Terms offered: Spring 2017, Spring 2014, Fall 2013
Meetings for the presentation of original work by faculty, visiting lecturers, and graduate students.

Integrative Biology Colloquium: Read More [+]

INTEGBI 296 Special Study for Graduate Students 1 - 4 Units

Terms offered: Spring 2016, Fall 2015, Spring 2015
Reading or other advanced study by arrangement with a staff member.

Special Study for Graduate Students: Read More [+]

INTEGBI 297 Directed Field Studies 1 - 8 Units

Terms offered: Spring 2017, Fall 2016, Spring 2016
Open to qualified students directly engaged in field studies.

Directed Field Studies: Read More [+]

INTEGBI 298 Special Study in Integrative Biology 1 - 12 Units

Terms offered: Fall 2018, Spring 2018, Fall 2017
Graduate research by small groups.

Special Study in Integrative Biology: Read More [+]

INTEGBI 299 Graduate Research 1 - 12 Units

Terms offered: Fall 2018, Spring 2018, Fall 2017
Credit awarded according to work planned and accomplished.

Graduate Research: Read More [+]

INTEGBI N299 Graduate Research 1 - 6 Units

Terms offered: Summer 2015 Second 6 Week Session, Summer 2010 10 Week Session, Summer 2007 10 Week Session
Graduate student research.

Graduate Research: Read More [+]

INTEGBI 304 Dissemination of Research: Your Interface with the Public 2 Units

Terms offered: Spring 2018, Spring 2017, Fall 2012
This course will consist of lectures and class discussions about mechanisms of communicating about science to the public. We will consider how to convey the issues, process, and findings of scientific research to a variety of audiences using different media (e.g., posters, web pages, newsletters, newspaper and magazine articles, books, television). Projects conducted by teams of students under the direct supervision of the instructors will
include preparation of outreach materials (e.g., posters, newsletters, web pages).
Dissemination of Research: Your Interface with the Public: Read More [+]

INTEGBI 305 Academic Survivorship 2 Units

Terms offered: Spring 2015, Fall 2011, Fall 2006
Series of lectures and workshops to prepare graduate students for many aspects of academic careers, including grant proposal writing, giving talks at meetings or to academic departments, preparing job applications and having job interviews, advising graduate students and postdocs, reviewing manuscripts and grant proposals, service activities and time management, working at teaching college vs. research universities, alternative careers, etc.

Academic Survivorship: Read More [+]

INTEGBI 375 Teaching Colloquium: Graduate Student Instructor Training 2 Units

Terms offered: Fall 2018, Fall 2017, Fall 2016
Series of workshops and seminars involving graduate students and faculty participation. The main objectives of this course are to train graduate students to become effective instructors and to discuss important issues that graduate students face when teaching undergraduate classes.

Teaching Colloquium: Graduate Student Instructor Training: Read More [+]

INTEGBI 400 Training in Stable Isotope Methods and Mass Spectrometry 1 Unit

Terms offered: Fall 2018, Fall 2017, Fall 2016
An intensive lecture and laboratory training course on the fundamental principles and practical applications of stable isotope methods in biogeochemistry, ecology, physiology, and environmental science. Topics covered are sample preparation, operating of an isotope ratio mass spectrometer, and analysis of stable isotope data. This course is required for all students interested in using the facilities housed in the Center for Stable Isotope Biogeochemistry
for their research.
Training in Stable Isotope Methods and Mass Spectrometry: Read More [+]

INTEGBI C407 Introduction to Scientific Diving 3 Units

Terms offered: Spring 2017, Spring 2016, Spring 2015
Diving physics, physiology, medicine, rescue, decompression, theory, navigation, environment, marine life, research methods, equipment, and University regulations. Course leads to University certification to use underwater life support apparatus for study or research under University auspices.

Introduction to Scientific Diving: Read More [+]

INTEGBI 601 Individual Study for Master's Students 1 - 8 Units

Terms offered: Spring 2016, Fall 2015, Spring 2015
Individual study for the comprehensive requirements in consultation with the major adviser. Units may not be used to meet either unit or residence requirements for a master's degree.

Individual Study for Master's Students: Read More [+]

INTEGBI 602 Individual Study for Doctoral Students 1 - 8 Units

Terms offered: Spring 2016, Fall 2015, Spring 2015
Individual study in consultation with the major adviser. Intended to provide an opportunity for qualified students to prepare themselves for the various examinations required for candidates for the Ph.D.

Individual Study for Doctoral Students: Read More [+]

INTEGBI N602 Individual Study for Doctoral Students 1 - 6 Units

Terms offered: Prior to 2007
Formerly < Paleon 602, Zoology 602, Botany 602, Physiol 602, Anatomy 602> Individual study in consultation with the major field adadviser. Intended to provide an opportunity for qualified students to prepare themselves for the various examinations required for candidates for the Ph.D.

Individual Study for Doctoral Students: Read More [+]

Faculty and Instructors

+ Indicates this faculty member is the recipient of the Distinguished Teaching Award.


David D. Ackerly, Professor. California biodiversity, climate change, adaptation.
Research Profile

Doris Bachtrog, Associate Professor. Evolution of sex and recombination, Y degeneration, dosage compensation, sexually antagonistic variation.
Research Profile

Bruce G. Baldwin, Professor. Biology, systematics and evolution of vascular plants, floristics, conservation biology, evolutionary processes, historical biogeography, evolutionary ecology.
Research Profile

Anthony D. Barnosky, Professor. Conservation biology, ecology, climate change, paleontology, paleobiology, paleoecology, evolution, macroecology, global change, mammals, extinction, biogeography.
Research Profile

George Bentley, Associate Professor. Hormones and behavior, neuroendocrinology of reproduction.
Research Profile

Jeffrey L. Boore, Adjunct Professor. Molecular biology, genetics, biology, conservation biology, genomics, evolution, genomes, DNA sequencing, systematics, population genetics, phylogeny.
Research Profile

Michael Robert John Boots, Professor.

+ Rauri C. K. Bowie, Associate Professor.

George A. Brooks, Professor. Exercise & environmental physiology, metabolism, lactate shuttle, crossover concept, lactate, lactic acidosis, traumatic brain injury, obesity, diabetes, the Warburg Effect in cancer, aging biology, isotope tracer technology, arterial-venous difference measurements, clamp technology, muscle biopsies, cell culture, mitochondrial biogenesis, energy substrate partitioning, carbohydrate metabolism, lipid metabolism, fatty acid reesterification, menopause.
Research Profile

Todd Dawson, Professor. Physiological plant ecology, evolutionary plant ecology, ecosystem processes, adaptations of plants, carbon, water, nitrogen.
Research Profile

Robert Dudley, Professor. Metabolism, biomechanics, butterflies, energetics, flight, gliding, hummingbirds, insects, paleophysiology.
Research Profile

Ivo Duijnstee, Assistant Adjunct Professor.

Michael B. Eisen, Professor. Genomics, genome sequencing, bioinformatics, animal development.
Research Profile

Paul V. A. Fine, Associate Professor. Speciation, plant ecology, plant evolutionary biology, floristics and phytogeography.
Research Profile

Seth Finnegan, Assistant Professor.

+ Robert J. Full, Professor. Energetics, comparative biomechanics, arthropod, adhesion, comparative physiology, locomotion, neuromechanics, biomimicry, biological inspiration, reptile, gecko, amphibian, robots, artificial muscles.
Research Profile

Oskar Hallatschek, Assistant Professor.

+ Tyrone Hayes, Professor. Genetics, amphibians, developmental endocrinology, steroid hormones, metamorphosis, sex differentiation, hormonal differentiation, African clawed frog, Japnanes Kajika, Pine Barrens treefrog.
Research Profile

Leslea Hlusko, Associate Professor. Primate evolution, paleontology, Ethiopia, Kenya, Tanzania, mammalian evolution, quantitative genetics, dental evolution, tooth development.
Research Profile

John P. Huelsenbeck, Professor. Computational biology, evolutionary biology, phylogenetics.
Research Profile

Daniela Kaufer, Associate Professor. Neuroscience, stress, neural stem cells, epilepsy, traumatic brain injury, blood brain barrier, prosocial behavior.
Research Profile

Mimi A. R. Koehl, Professor. Biomechanics, insects, invertebrate functional morphology, fluid dynamics, aerodynamics, marine animals, filtration, gliding vertebrates.
Research Profile

Britt Koskella, Assistant Professor.

+ Eileen A. Lacey, Associate Professor. Evolutionary biology, population, mammals, behavioral ecology, vertebrates, molecular genetics, subterranean rodents, Argentina, Chile.
Research Profile

Cindy Looy, Assistant Professor. Paleoecology, paleobotany, palynology.
Research Profile

Charles Marshall, Professor.

Jimmy Mcguire, Associate Professor. Historical biogeography, evolutionary biology, Southeast Asia, population genetics, hummingbirds, functional morphology, vertebrate systematics, phylogenetic analysis, life history evolution, Bayesian methods, Southeast Asian flying lizards.
Research Profile

Brent D. Mishler, Professor. Evolutionary biology, development, ecology, systematics, phylogeny, comparative genomics, green plants, bryophytes, mosses, reproductive biology, phylogenetics, chloroplast DNA, classification, species concepts, biodiversity, Darwin.
Research Profile

Michael Nachman, Professor. Population genetics, evolution, genomics, mammalian evolution.
Research Profile

Rasmus Nielsen, Professor. Statistical and computational aspects of evolutionary theory and genetics.
Research Profile

Kevin Padian, Professor. Evolutionary biology, paleontology, systematics, functional morphology, Mesozoic vertebrate paleontology, terrestrial vertebrate fauna, pterosaurs, Mesozoic era, bones of extinct reptiles.
Research Profile

Nipam Patel, Professor. Genetics, evolution, crustaceans, insects, arthropods, homeotic (Hox) genes, segmentation, embryonic pattern formation, neural patterning.
Research Profile

Mary E. Power, Professor. Freshwater ecology, food webs, trophic dynamics, northern California rivers, watersheds.
Research Profile

Carl Rothfels, Assistant Professor.

Michael Shapira, Assistant Professor in Residence. Molecular biology, aging, host-pathogen interactions.
Research Profile

Ellen L. Simms, Professor. Evolution, plant ecology, microbial ecology, ecological genetics, symbiosis, herbivores, pathogens, mutualists.
Research Profile

Wayne Sousa, Professor. Community ecology, estuarine host-parasite interactions, canopy gaps, Caribbean coast of Panama.
Research Profile

Chelsea Specht, Associate Professor. Molecular evolution, adaptation, plant systematics, evolution of development (evo-devo), evolution of form and function, plant morphology, comparative phylogenetics, floral developmental evolution, diversification rates.
Research Profile

Jonathon Stillman, Assistant Adjunct Professor. Marine environmental physiology, global change biology, functional genomics.
Research Profile

Tim White, Professor. Anthropology, Africa, paleontology, paleoanthropology, human evolution, human osteology, taphonomy, zooarchaeology, cannibalism, American Southwest, Ethiopia.
Research Profile

Caroline Margaret Williams, Assistant Professor. Evolution, physiology, ecophysiology, metabolism, insect, winter, adaptation, thermal biology.
Research Profile


Kelly Agnew, Lecturer.

Thomas J. Carlson, Lecturer SOE. Molecular biology, ethnobotany, Africa, North America, ecology, medicine, systematics, evolution of human disease, ethnoecology, ethnoepidemiology, Asia, Pacific Islands, South America, nutritional ethnobotany, pharmacology, ecosystem management.
Research Profile

Christopher Hobbs, Lecturer.

Stephen Lew, Lecturer.

Stefania Mambelli, Lecturer.

Lisa A. Margerum, Lecturer.

Tim Markowitz, Lecturer.

Michael S. Park, Lecturer.

Helian Joel Ratsirarson, Lecturer.

Andrew C. Rush, Lecturer.

Christopher Schmitt, Lecturer.

Emeritus Faculty

Roy L. Caldwell, Professor Emeritus. Ecology, evolution, Invertebrates, animal behavior, behavioral ecology, marine biology, stomatopods, crustaceans, cephalopods, octopus, mating systems, communication, sensory ecology, aggressive behavior, coral reef restoration.
Research Profile

William A. Clemens, Professor Emeritus. Evolution of mesozoic, cenozoic terrestrial biotas, mesozoic mammals, phylogenetic interrelationships, locomotor evolution, evolutionary diversification.
Research Profile

+ Marian C. Diamond, Professor Emeritus. Environment, neuroanatomy, immune functions, hormones, mammalian forebrain structures, Cambodian orphanage, cerebral neocortex.
Research Profile

Carole S. Hickman, Professor Emeritus. Systematics, evolutionary paleobiology, morphology, molluscs, macroevolutionary trends and patterns, Cenozoic Era, fossil record, evolutionary history and structure.
Research Profile

Steven L. Lehman, Professor Emeritus. Motor control, stroke patients, cells, muscles, muscle fatigue, repetitive motion disorders.
Research Profile

Paul Licht, Professor Emeritus. Steroid hormones, comparative endocrinology, endocrine system, steroidal metabolism of tissues, thyroid, vitamin D, binding proteins.
Research Profile

William Z. Lidicker, Professor Emeritus. Conservation biology, ecology, mammalogy.
Research Profile

David R. Lindberg, Professor Emeritus. Developmental biology, evolutionary biology, ecology, comparative morphology, phylogenetic studies, molecular techniques, patellogastropoda, evolution in patellacean faunas, California land snail taxa, gastropoda.
Research Profile

Jere Lipps, Professor Emeritus. Ecology, paleontology, molecular phylogenetics, evolution of marine biotas, paleoenvironments, coral reef, Antarctic habitats, seismic histories, astrobiology, Charles Darwin.
Research Profile

Charles S. Nicoll, Professor Emeritus. Cancer, mammals, hormones, growth regulation, diabetic subjects, insulin, reproductive cycles of women, non-human primates.
Research Profile

+ James L. Patton, Professor Emeritus. Evolutionary biology, North America, biogeography, South America, morphology, mammalogy, phylogenesis, speciation, populations, genome structure, molecular divergence, gopher population dynamics.
Research Profile

Thomas M. (Zack) Powell, Professor Emeritus. Aquatic ecology, fish, Invertebrates, oceanography, lakes, estuaries, ocean, planktonic ecosystems, climate, remote sensing.
Research Profile

Thelma Rowell, Professor Emeritus.

Rudolf Schmid, Professor Emeritus. Botany, plant science.
Research Profile

Montgomery Slatkin, Professor Emeritus. Evolutionary theory, genetic evolution, natural populations of plants and animals populations, human populations, natural selection structure genomes.
Research Profile

Glenys Thomson, Professor Emeritus.

James Valentine, Professor Emeritus.

David Wake, Professor Emeritus. Evolutionary and conservation biology.
Research Profile

Marvalee H. Wake, Professor Emeritus. Development, evolution, systematics, amphibians, reproductive biology, vertebrate evolutionary morphology, fishes, reptiles, comparative analysis, biodiversity issues.
Research Profile

Donald P. Weston, Professor Emeritus. Pesticides, invertebrate ecology, ecotoxicology, pollutant exposure, bioaccumulation, management of contaminated sediments, environmental effects of antibiotics, deposit-feeding invertebrates.
Research Profile

Contact Information

Department of Integrative Biology

3060 Valley Life Sciences Building

Phone: 510-642-5024

Fax: 510-643-6264

Visit Department Website

Department Chair

Robert Dudley, PhD

5018 Valley Life Sciences Building

Phone: 510-642-1555

Head Graduate Adviser

Brent Mishler

Graduate Affairs Office

299 Life Sciences Addition

Phone: 510-643-7330

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