Courses
This is a comprehensive listing of courses taught within EEB. With the exception of major requirements, not all courses are guaranteed to be taught in the future. Please consult the Course Search (link is external) for a listing of courses offered this semester.
LINKS: Undergraduate Courses | Graduate Courses | Course Search (link is external)
Biochemistry and Biophysics (BIOL 1010)
The study of life at the molecular level. Topics include the three-dimensional structures and function of large biological molecules, the human genome, and the design of antiviral drugs to treat HIV/AIDS.
The first of four modules in a yearlong foundational biology sequence; meets for the first half of the term.
Cell Bio & Membrane Physiology (BIOL 1020)
The study of cell biology and membrane physiology. Topics include organization and functional properties of biological membranes, membrane physiology and signaling, rough endoplasmic reticulum and synthesis of membrane/secretory membrane proteins, endocytosis, the cytoskeleton, and cell division.
The second of four modules in a yearlong foundational biology sequence; meets for the second half of the term. Prerequisite: BIOL 1010.
Genes and Development (BIOL 1030)
Foundation principles for the study of genes, genetics, and developmental biology. How genes control development and disease; Mendel’s rules; examples of organ physiology.
The third of four modules in a yearlong foundational biology sequence; meets for the first half of the term. Prerequisites: BIOL 1010, 1020
Ecology & Evolutionary Biology (BIOL 1040)
The study of evolutionary biology, animal behavior, and the history of life. Evolutionary transitions and natural selection. Adaptation at genic, chromosomal, cellular, organismal, and supra-organismal levels. Distributional and social consequences of particular suites of organismal adaptations.
The fourth of four modules in a yearlong foundational biology sequence; meets for the second half of the term. Prerequisites: BIOL 1010, 1020, and 1030.
The Ecology of Food (EEB 0035)
Food and ecology are inextricably linked, both in producing domesticated food through agriculture and livestock and in harvesting wild plants and animals. Furthermore, the production and consumption of food have downstream consequences through energy consumption, food waste, trophic interactions, and the transportation of food around the globe. These topics link to many of the fundamental concepts of ecology, including population biology, the niche, trophic interactions, nitrogen cycling, and the effects on biodiversity. In this class, we explore these topics intensively through a combination of lectures, readings, and interactive field trips to on/near campus ecosystems, including the Marsh Botanical Gardens, the Yale Sustainable Farm, a nearby forest and salt marsh, and an orchard. Each week, we meet twice for 50 minutes for a combination of lecture and discussion and for 110 minutes for field trips, discussions, and guest lectures.
Enrollment limited to first-year students.
1 credit for Yale College students
Collections of the Peabody Museum (EEB 0040)
The Evolution of Beauty (EEB 0050)
Exploration of the fundamental mechanism of sexual selection and mate choice, and the patterns of display trait and mating preference coevolution. Additional topics include what happens when the freedom of mate choice is infringed or disrupted by sexual coercion or sexual violence; and the role of aesthetic evolution and sexual conflict in the evolution of human sexuality, pleasure, and sexual diversity. Enrollment limited to first-year students.
Preregistration required; see under Frist Year Seminar Program.
1 Yale College course credit(s).
Biology of Malaria, Lyme and Other Vector Borne Disease (EEB 1106 )
Introduction to the biology of pathogen transmission from one organism to another by insects; special focus on malaria, dengue, and Lyme disease. Biology of the pathogens including modes of transmission, establishment of infection, and immune responses; the challenges associated with vector control, prevention, development of vaccines, and treatments. Intended for non–science majors; preference to freshmen and sophomores.
Prerequisite: high school biology.
1 Yale College course credit(s)
Conservation Biology (EEB 1115)
An introduction to ecological and evolutionary principles underpinning efforts to conserve Earth’s biodiversity. Efforts to halt the rapid increase in disappearance of both plants and animals. Discussion of sociological and economic issues.
1 Yale College course credit(s)
History of Life (EEB 1125)
Examination of fossil and geologic evidence pertaining to the origin, evolution, and history of life on Earth. Emphasis on major events in the history of life, on what the fossil record reveals about the evolutionary process, on the diversity of ancient and living organisms, and on the evolutionary impact of Earth’s changing environment.
1 Yale College course credit(s)
Plants and People (EEB 1145)
The interaction of plants and people throughout history explored from biological, historical, anthropological, and artistic perspectives. Basic botany; plants in the context of agriculture; plants as instruments of trade and societal change; plants as inspiration; plants in the environment. Includes field trips to the greenhouses at Yale Marsh Botanical Garden, the Yale Peabody Museum and Herbarium, the Yale Farm, and the Yale Art Gallery.
1 Yale College course credit(s)
Virus Discovery and Evolution Lab (EEB 1175L)
An inquiry-based, hands-on introduction to sampling bacteriophages (bacteria-specific viruses) from natural environments. Emphasis on lab methods to characterize viruses via growth assays and genome sequencing, and to experimentally evolve viruses on bacteria. Readings and discussion on virus biodiversity, role of viruses in the environment, and virus applications to solve human problems.
Introduction to Statistics: Life Sciences (EEB 2210)
Statistical and probabilistic analysis of biological problems, presented with a unified foundation in basic statistical theory. Problems are drawn from genetics, ecology, epidemiology, and bioinformatics.
1 Yale College course credit(s)
Data Science for Ecology & Conservation (EEB 2212)
The course provides a practical foundation for data-driven inference and prediction in biodiversity science. The first part of the course is focused on providing a unified foundation for statistical analysis in ecology and aims to complement more general introductory statistical courses. Specific foci include an introduction and practice around General Linear Models (GLMs), and their use for hypothesis testing, prediction, and forecasting, as well as maximum likelihood and non-parametric approaches, all focused on biodiversity. The course then examines the promise and practice of emerging machine learning approaches specifically for prediction. With these foundations in place, the course will tackle several case studies addressing biogeography, community ecology, local conservation, and large-scale conservation decision-making. Students will be expected to work through cases studies individually or in small groups and present on their findings in class. The course will familiarize students with R, the leading analysis and visualization software in much of the life sciences, through in-class guidance, homework and problem sets.
The course assumes that students are familiar with the core concepts of probability and statistical analysis and have at minimum completed an introductory statistics course such as S&DS 100 (link is external) or similar by approval.
General Ecology (EEB 2220/5220)
The theory and practice of ecology, including the ecology of individuals, population dynamics and regulation, community structure, ecosystem function, and ecological interactions at broad spatial and temporal scales. Topics such as climate change, fisheries management, and infectious diseases are placed in an ecological context.
Prerequisite: MATH 112 or equivalent.
1 Yale College course credit(s)
Laboratory for Principles of Ecology, Evolutionary Biology, and the Tree of Life (EEB 2223L/5223L)
Study of evolutionary novelties, their functional morphology, and their role in the diversity of life. Introduction to techniques used for studying the diversity of animal body plans. Evolutionary innovations that have allowed groups of organisms to increase their diversity.
0.5 Yale College course credit(s)
Evolutionary Biology (EEB 2225/5225)
An overview of evolutionary biology as the discipline uniting all of the life sciences. Reading and discussion of scientific papers to explore the dynamic aspects of evolutionary biology. Principles of population genetics, paleontology, and systematics; application of evolutionary thinking in disciplines such as developmental biology, ecology, microbiology, molecular biology, and human medicine.
1 Yale College course credit(s)
Ecology and Evolution of Infectious Disease (EEB 2228/5228)
Overview of the ecology and evolution of pathogens (bacteria, viruses, protozoa) and their impact on host populations. Topics include theoretical concepts, ecological and evolutionary dynamics, molecular biology, and epidemiology of ancient and emerging diseases.
1 Yale College course credit(s)
Field Ecology (EEB 2230/5230)
A field-based introduction to ecological research, using experimental and descriptive approaches, comparative analysis, and modeling for field and small-group projects. Weekly field trips explore local lake, salt marsh, rocky intertidal, traprock ridge, and upland forest ecosystems. Includes one Saturday field trip and a three-day trip during the October recess. Concurrently with or after EEB 2220 or with permission of instructor.
1 Yale College course credit(s)
Behavioral Ecology (EEB 2242/5242)
An introduction to the study of animal behavior from an evolutionary and ecological perspective. Topics include decision-making, group living and cooperation, sexual selection and mating behavior, signaling and communication. In addition to lectures, in-class discussions and activities, students engage in the material by design and implement their own research projects.
Prerequisite: BIOL 104 (link is external), or permission of instructor.
Plant Diversity & Evolution (EEB 2246/5246)
This course has several, interrelated objectives. First, it serves as an introduction to the science of phylogenetics, providing an overview of both the theory and methodology involved in constructing phylogenetic trees, and how to use trees to study character and organismal evolution. For our second objective, we put this new framework to immediate use by using phylogeny to explore and illustrate 400 million years of land plant evolution, with emphasis on the diversity of flowering plants. The course examines major trends in plant evolution from functional, ecological, and bio-geographical perspectives. Students acquire a basic understanding of 1) phylogenetic approaches to comparative biology, 2) plant anatomy and morphology, 3) evolutionary relationships among the major land plant clades (with emphasis on the flowering plants), and 4) major evolutionary trends that have significantly shaped the diversity of plant life that we see today. The third and most important objective is to instill in students the ability to look at any biological problem through the lens of “phylogeny-colored glasses”- a powerful way to examine the complexity of life that surrounds (and includes!) us.
Laboratory for Plant Diversity & Evolution (EEB 2247L/5247L)
Hands-on experience with the plant groups examined in the accompanying lectures. Local field trips.
To be taken concurrently with EEB 2246.
0.5 Yale College course credit(s)
Biology Of Terrestrial Arthropods (EEB 2250/5250)
Evolutionary history and diversity of terrestrial arthropods (body plan, phylogenetic relationships, fossil record); physiology and functional morphology (water relations, thermoregulation, energetics of flying and singing); reproduction (biology of reproduction, life cycles, metamorphosis, parental care); behavior (migration, communication, mating systems, evolution of sociality); ecology (parasitism, mutualism, predator-prey interactions, competition, plant-insect interactions).
To be taken concurrently with EEB 2251L.
1 Yale College course credit(s)
Laboratory for Biology Of Terrestrial Arthropods (EEB 2251L/5251L)
Comparative anatomy, dissections, identification, and classification of terrestrial arthropods; specimen collection; field trips.
Concurrently with or after EEB 2250.
0.5 Yale College course credit(s)
Invertebrates (EEB 2255/5255)
An overview of animal diversity that explores themes including animal phylogenetics (evolutionary relationships), comparative studies of evolutionary patterns across species, organism structure and function, and the interaction of organisms with their environments. Most animal lineages are marine invertebrates, so marine invertebrates are the focus of most of the course.
To be taken concurrently with EEB 2256L.
1 Yale College course credit(s)
Laboratory for Invertebrates (EEB 2256L/5256L)
The study of invertebrate anatomy and diversity in a laboratory and field setting. Activities will include will examine live animals and museum specimens, as well as local field trips. Some field trips will fall on weekends.
This lab must be taken concurrently with the lecture EEB 2255.
0.5 Yale College course credit(s)
The Biology of Sharks and Their Relatives (EEB 2262)
An integrative course that examines the biology of sharks and other cartilaginous fishes (Class Chondrichthyes) from a variety of perspectives. Students learn about the taxonomy and systematics, paleontology, functional anatomy, behavior, physiology, ecology, and cultural significance of sharks. Coursework includes answers to discussion prompts, guided review of scientific literature, and in-class exams that allow students to demonstrate their understanding of chondrichthyan biology and sharks’ unique place in the vertebrate tree of life. To be taken with EEB 2263L (link is external).
The Biology of Sharks and Their Relatives Laboratory (EEB 2263L)
This is a hands-on, specimen-based overview of the fossil record, comparative anatomy, functional morphology, and biodiversity of sharks and their relatives, the skates, rays, and ratfish. Students examine and dissect fresh and preserved specimens and use the fossil remains of extinct sharks to investigate the evolution of cartilaginous fishes. This course should be taken concurrently with EEB 2262 (link is external), The Biology of Sharks and Their Relatives.
Ichthyology (EEB 2264/5264)
A survey of fish diversity, including jawless vertebrates, chimaeras and sharks, lungfishes, and ray-finned fishes. Topics include the evolutionary origin of vertebrates, the fossil record of fishes, evolutionary diversification of major extant fish lineages, biogeography, ecology, and reproductive strategies of fishes
1 Yale College course credit(s)
Laboratory for Ichthyology (EEB 2265L/5265L)
Laboratory and field studies of fish diversity, form, function, behavior, and classification. The course primarily involves study of museum specimens and of living and fossil fishes. Concurrently with EEB 2264.
0.5 Yale College course credit(s)
Bird Behavior (EEB 2269)
A seminar discussion of classic and recent scientific literature on topics in bird behavior. Students develop experience in critical reading of the literature through the exploration of topics in bird behavior including courtship, breeding behavior, song and song learning, foraging ecology, migration and orientation, and sensory ecology.
Prerequisite: BIOL 1040 (link is external) or permission of the instructor.
Ornithology (EEB 2272/5272)
An overview of avian biology and evolution, including the structure, function, behavior, and diversity of birds. The evolutionary origin of birds, avian phylogeny, anatomy, physiology, neurobiology, breeding systems, and biogeography.
Enrollment limited to 50.
1 Yale College course credit(s)
Laboratory for Ornithology (EEB 2273L/5273L)
Laboratory and field studies of avian morphology, diversity, phylogeny, classification, identification, and behavior.
Enrollment limited to 12.
0.5 Yale College course credit(s)
Biological Oceanography (EEB 2275/EVST 3400)
Exploration of a range of coastal and pelagic ecosystems. Relationships between biological systems and the physical processes that control the movements of water and productivity of marine systems. Anthropogenic impacts on oceans, such as the effects of fishing and climate change. Includes three Friday field trips.
Enrollment limited to 15.
1 Yale College course credit(s)
Evolution and Ecology of Marine Vertebrates (EEB 2280)
Fishes, reptiles, and mammals all include species that spend all or much of their lives in or near the ocean. In this course, we survey those taxa, paying particular attention to how the biotic and abiotic aspects of marine environments have shaped organismal morphology, physiology, and behavior. Students learn about the biology of marine fishes as well as extinct marine reptiles, modern sea turtles and sea snakes, shore birds, and whales and their relatives. This course includes discussions of the conservation challenges facing each group. Because of our location and the deep historical connection our region has to fisheries, including the whaling industry of the eighteenth and nineteenth centuries, we emphasize the marine vertebrates of New England and the Mid-Atlantic coasts. This course includes a mandatory field trip to a regional aquarium or local shoreline.
Students are strongly encouraged to have taken the introductory biology sequence (BIOL 101 to 104) or have the equivalent understanding of biology, evolution, and ecology.
Comparative Developmental Anatomy of Vertebrates (EEB 2290)
A survey of the development, structure, and evolution of major vertebrate groups. Topics include the micro-anatomy of major organ systems, the developmental underpinnings of the vertebrate body plan, and the development, structure and evolution of the major organ systems such as the locomotory system, sensory organs, digestive tract, reproductive tract, and nervous system.
To be taken concurrently with EEB 2291L (link is external).
1 Yale College course credit(s)
Laboratory for Comparative Anatomy of Vertebrates (EEB 2291L)
Microscopic examination of histological and embryological preparations. Dissection of selected vertebrate species including shark, bony fish, frog, lizard, and rat.
To be taken with EEB 2290 (link is external).
0.5 Yale College course credit(s)
Life in Motion: Ecological and Evolutionary Physiology (EEB 2295)
Physiology is the study of the functions that organisms perform and how they use those functions to interact with the environment. To survive, grow, and reproduce, all organisms must acquire energy and avoid conditions that exceed their physiological limits. These interactions all involve motion—ions traveling across membranes, muscle fibers twitching, respiration, and locomotion, to name a few. In this course, we tackle physiological processes from both “bottom up” and “top down” approaches, with integration among these dimensions, to extract general physiological rules of life. Then, we link our discoveries to the broader context of ongoing global change, and consider whether and how organisms can physiologically respond to contemporary selective pressures. While the course focuses heavily on animal physiology, plants, fungi, and microbes are also featured.
Prerequisites: BIOL 1010, 1020, 1030, 1040, and CHEM 1610, or permission of the instructor.
1 Yale College course credit(s)
Plant Ecology (EEB 305/705)
The study of plant interactions with their environment, at the level of individuals, and of how plant-plant interactions mediate environmental interactions at the level of populations, communities, and ecosystems. Incorporation of empirical and theoretical perspectives, emphasizing the empirical origins of concepts in plant ecology and effective empirical tests of conceptual and mathematical predictions.
1 Yale College course credit(s)
Plant ecology is the study of plant interactions with their environment, at the level of individuals, and of how plant-plant interactions mediate environmental interactions at the level of populations, communities, and ecosystems. The course incorporates empirical and theoretical perspectives, emphasizing the empirical origins of concepts in plant ecology and effective empirical tests of conceptual and mathematical predictions. Students read the primary scientific literature extensively, both for content and to build familiarity with methodological standards and the scientific writing.
1 Yale College course credit(s).
Plant ecology is the study of plant interactions with their environment, at the level of individuals, and of how plant-plant interactions mediate environmental interactions at the level of populations, communities, and ecosystems. The course incorporates empirical and theoretical perspectives, emphasizing the empirical origins of concepts in plant ecology and effective empirical tests of conceptual and mathematical predictions. Students read the primary scientific literature extensively, both for content and to build familiarity with methodological standards and the scientific writing.
Primate Behavior and Ecology (EEB 3300)
Socioecology of primates compared with that of other mammals, emphasizing both general principles and unique primate characteristics. Topics include life-history strategies, feeding ecology, mating systems, and ecological influences on social organization.
Community Ecology (EEB 3320/5320)
This course covers core questions in community ecology related to species interactions, species coexistence theory, species-environment interactions, the consequences of biological diversity, spatial ecology, food webs, and eco-evolutionary interactions. Lectures emphasize the theoretical and conceptual foundations of these topics and incorporate the empirical and experimental evidence supporting and confronting contemporary views.
Prerequisites: EEB 2220 (link is external) or EVST 2200 (link is external), or with permission of instructor.
1 Yale College course credit(s).
Philosophy of Biology (EEB 3321)
This course is an introduction to the philosophy of biology, with application to specific current problems. It focuses on two major strands of thinking seeking answers to fundamental and complementary questions: “How do we know?” (epistemology) and “What things really exist in the world?” (ontology). These two themes have the most important impact on the practice of science, as they pertain to the nature of the scientific enterprise and how it works (epistemology and philosophy of science), as well as what scientists consider part of reality (=science related ontology: unicorns and phlogiston NO; atoms, electrons, YES; but what about species and genes? Do they have the same status as atoms?). In each of these fields of philosophy we outline the main positions and then discuss how they apply to past and current debates in biology, in particular, but not exclusively, evolutionary biology.
Prerequisite: a semester of biology or a semester of philosophy.
1 Yale College course credit(s)
An introduction to the philosophy of biology, with application to specific current problems. The course focuses on two major strands of thinking seeking answers to two fundamental and to some extent complementary questions: “How do we know?” [epistemology] and “What things really exist in the world?” [ontology]. These two themes have the most important impact on the practice of science, as they pertain to the nature of the scientific enterprise and how it works [epistemology and philosophy of science], as well as what scientists consider part of reality [science-related ontology: unicorns and phlogiston, NO; atoms, electrons, YES; but what about species and genes? Do they have the same status as atoms?]. In each of these fields of philosophy we outline the main positions and discuss how they apply to past and current debates in biology—in particular, but not exclusively, evolutionary biology.
1 Yale College course credit(s)
Evolutionary Genetics (EEB 3322/5322)
Genetic variation is the currency by which natural selection is translated into evolutionary change. In this course we dissect patterns of genetic variation using an evolutionary mindset to ultimately understand what shapes genetic variation in nature and the potential for species to adapt to new and changing environments. This class unites two foundational fields of evolutionary genetics; quantitative genetics (the study of the genetic basis of complex traits) and population genetics (the study of gene variant frequencies across time and space), with an ultimate goal of understanding evolutionary change in nature. Although this course is lecture based, there is much opportunity for hands-on learning. Students use real-life and simulated genetic data to map the genetic basis of traits and investigate the evolutionary forces responsible for shaping genetic variation in nature. We also discuss how quantitative and population genetics theory are applied to the modern genomic era, particularly in the context of detecting genomic signatures of adaptation. Lastly, we discuss the application of evolutionary genetics to human populations, including the usefulness and missteps of these applications for science and society.
Prerequisite: EEB 2225 (link is external), Evolutionary Biology.
Limnology (EEB 3325)
Limnology, the study of the physical, chemical, and biological properties of inland waters, focuses on lakes where physical (light, temperature, and mixing) and chemical (dissolved elements and compounds) properties interact with the ecology and evolution of organisms. Topics include origins and morphology of inland waters; physical and chemical properties; diversity and interactions among the organisms found in lakes; historical perspectives; and understanding conservation and management in the context of global change. Frequent field trips to local freshwater ecosystems.
Prerequisites: EEB 2220 and EEB 2225, or with permission of instructor.
1 Yale College course credit(s).
Limnology, the study of the physical, chemical, and biological properties of inland waters, focuses on lakes where physical (light, temperature, and mixing) and chemical (dissolved elements and compounds) properties interact with the ecology and evolution of organisms. Topics include origins and morphology of inland waters; physical and chemical properties; diversity and interactions among the organisms found in lakes; historical perspectives; and understanding conservation and management in the context of global change. Frequent field trips to local freshwater ecosystems.
Plant Structure and Function (EEB 3326)
This is an advanced botany course, preferably for students that have taken EEB 2246 in addition to BIO 1040; otherwise permission must be obtained from the instructor. A keen interest in plants is a must.
1 Yale College course credit(s).
Plant Structure Laboratory (EEB 3327L)
Laboratory for Plant Structure and Function (EEB 3327L)
This is an advanced botany course, preferably for students that have taken EEB 2246 in addition to BIO 1040; otherwise permission must be obtained from the instructor. A keen interest in plants is a must.
1 Yale College course credit(s).
Ecosystem Ecology (EEB 3330/5330)
Ecosystem ecology asks how abiotic and biotic processes come together to shape the diversity in form and function across Earth’s ecosystems, from the flow of energy and materials through the environment, to how communities of organisms interact with their environment. This course examines the factors that influence ecosystem structure and function: the processes that shape how energy, water, carbon, and nutrients cycle through ecosystems, the role of disturbance on these processes, and feedbacks from human-induced global change.
Prerequisites: EEB 2220(link is external) (link is external) or EVST 2220(link is external) (link is external), or with permission of instructor.
1 Yale College course credit(s).
Evolution and Medicine (EEB 3335/5335)
Introduction to the ways in which evolutionary science informs medical research and clinical practice. Diseases of civilization and their relation to humans’ evolutionary past; the evolution of human defense mechanisms; antibiotic resistance and virulence in pathogens; cancer as an evolutionary process. Students view course lectures on line; class time focuses on discussion of lecture topics and research papers.
Prerequisite: BIOL 101–01040.
1 Yale College course credit(s)
Culture and Human Evolution (EEB 3336)
Examination of the origins of human modernity in the light of evolutionary and archaeological evidence. Understanding, through a merger of evolutionary reasoning with humanistic theory, the impact of human culture on natural selection across the last 250,000 years.
1 Yale College course credit(s)
Microbial Ecology (EEB 3340/5340)
When thinking about microbes what comes to mind are usually diseases and unpleasant smells from the fridge or the basement. Nevertheless, microbes and the communities they form are key contributors to our wellbeing and the functioning of the planet. This course provides an introduction to microbial ecology, with an emphasis on how microbial systems differ from their macroscopic counterparts, including defining a microbial species; sampling/experimenting with microbes; principles of microbial growth, metabolism, and death; species interactions and community assembly in different environments; microbial community functions; elements of microbial evolution.
BIOL 1010 (link is external), BIOL 1020 (link is external), BIOL 1030 (link is external), & BIOL 1040 (link is external). General Ecology EEB 2220 (link is external) and MCDB 2900 (link is external) are encouraged but not required.
1 credit for Yale College students
Primate Diversity and Evolution (EEB 3342/5342)
The diversity and evolutionary history of living and extinct primates. Focus on major controversies in primate systematics and evolution, including the origins and relationships of several groups. Consideration of both morphological and molecular studies. Morphological diversity and adaptations explored through museum specimens and fossil casts.
Requires Permission of the Instructor
Recommended preparation: ANTH 1400 (link is external).
I Yale College course credit(s).
Ecosystem Measurements for Ecology and Conservation (EEB 3343L)
This course familiarizes students with how ecology is used on the ground for conservation. It is structured in two parts: The first part of the class will be dedicated to active hands-on learning where students obtain formal training in broad range of field and lab methods and analyses used in ecological field research. Topics covered include carbon stock measurement, biodiversity assessment, utilization of digital conservation resources, experimental design, sampling methodology, and statistical analysis. The course includes intensive field exercises focused on forest measurements and soil analysis that ecologists use to understand ecosystem function. The second component of the course allows students to use these skills to design, conduct, analyze and present data in the form of a rapid ecological assessment or group research project on a local property of conservation importance.
Prerequisites: BIOL 104 (link is external) or instructor permission
Evolutionary Theory (EEB 3352/5352)
This course introduces students to the theory behind evolutionary biology. The aim of the course is for the student to understand how evolution works, focusing on the quantitative and predictive theory that is the backbone of modern evolutionary thinking. The course covers three main areas: An introduction to population genetics, an introduction to quantitative genetics and the genotype-phenotype map, and an introduction to life-history evolution. To master this material and to put the concepts studied in class into practice, students work on weekly problem sets. Through the completion of the course assignments, students gain valuable quantitative and mathematical modeling skills.
Prerequisites: One of the following: EEB 2225 (link is external), PHYS 1700 (link is external)/1710 (link is external) or 1800 (link is external)/1810 (link is external) or permission of the instructor.
Phylogenetic Biology (EEB 3354/5354)
Phylogenetic Biology is the study of the evolutionary relationships between organisms, and the use of evolutionary relationships to understand other aspects of organism biology. This course surveys phylogenetic methods, providing a detailed picture of the statistical, mathematical, and computational tools for building phylogenies and using them to study evolution. We also examine the application of these tools to particular problems in the literature and emerging areas of study.
Prerequisites: EEB 2225 and an organismal course.
1 Yale College course credit(s)
Tropical Field Biology (EEB 3362)
Firsthand experience of a region can inspire great insights and understanding of ecology and evolutionary biology. This course immerses students in the communities and ecosystems of a single tropical region each year, but locations rotate among a small group of sites. We spend the first half of the semester learning about the geology, history, biomes and organisms of the region.
Prerequisites: EEB 2220 (link is external), EEB 2225 (link is external), and permission of the instructor.
Topics in Vertebrate Ecomorphology (EEB 3375)
Ecomorphology is a field that bridges ecology and evolutionary biology. Researchers studying organisms’ ecomorphology ask questions like, “What does the morphology of an organism tell us about its relationship with its environment” and “How are correlations between morphology and ecology influenced by behavior?” The answers to questions like these inform evolutionary hypotheses based on natural selection and help to explain the amazing diversity of life forms that surround us. In this course, we explore the links between organismal form, function, ecology, and evolution using a series of readings and guided discussions. Students also learn many of the fundamentals associated with crafting and revising publishable scientific writing–a must for those seeking research-based graduate education in the sciences. By the end of the semester, students refine their critical thinking and scientific writing skills, and they have a newfound awareness of one of the most integrative and fascinating branches of vertebrate biology.
1 credit for Yale College students
Life History Evolution (EEB 3380)
Life history evolution studies how the phenotypic traits directly involved in reproductive success are shaped by evolution to solve ecological problems. The intimate interplay between evolution and ecology.
Requires Permission of the Instructor
After EEB 2220 and 2225, or with permission of instructor.
1 Yale College course credit(s).
Science of Complex Systems (EEB 4428)
Introduction to the quantitative analysis of systems with many degrees of freedom. Fundamental components in the science of complex systems, including how to simulate complex systems, how to analyze model behaviors, and how to validate models using observations. Topics include cellular automata, bifurcation theory, deterministic chaos, self-organized criticality, renormalization, and inverse theory.
Prerequisite: PHYS 4000, MATH 2470, or equivalent.
1 Yale College course credit(s)
Human Osteology (EEB 4464)
A lecture and laboratory course focusing on the characteristics of the human skeleton and its use in studies of functional morphology, paleodemography, and paleopathology. Laboratories familiarize students with skeletal parts; lectures focus on the nature of bone tissue, its biomechanical modification, sexing, aging, and interpretation of lesions.
1 Yale College course credit(s)
Tutorial (EEB 4469)
Individual or small-group study for qualified students who wish to investigate an area of ecology or evolutionary biology not presently covered by regular courses. A student must be sponsored by a faculty member who sets requirements and meets weekly with the student. One or more written examinations and/or a term paper are required. To register, the student must submit a written plan of study approved by the faculty instructor to the director of undergraduate studies. Students are encouraged to apply during the term preceding the tutorial. Proposals must be submitted no later than the first day of the second week of the term in which the student enrolls in the tutorial. The final paper is due in the hands of the director of undergraduate studies by the last day of reading period in the term of enrollment. In special cases, with approval of the director of undergraduate studies, this course may be elected for more than one term, but only one term may be counted as an elective toward the requirements of the major. Normally, faculty sponsors must be members of the EEB department.
Requires Permission of the Instructor
http://eeb.yale.edu/sites/default/files/eeb_469_tutorial_form_rev_8.24.1…
1 Yale College course credit(s).
Senior Tutorial (EEB 4470)
Tutorial for seniors in the B.A. degree program who elect a term of independent study to complete the senior requirement. A student must be sponsored by a faculty member who sets requirements and meets weekly with the student. One or more written examinations and/or a term paper are required. To register, the student must submit a written plan of study approved by the faculty instructor to the director of undergraduate studies. Students are encouraged to apply during the term preceding the tutorial. Proposals must be submitted no later than the first day of the second week of the term in which the student enrolls in the tutorial. The final paper is due in the hands of the director of undergraduate studies by the last day of reading period in the term of enrollment. Normally, faculty sponsors must be members of the EEB department.
Requires Permission of the Instructor
Enrollment limited to seniors. Fulfills the senior requirement for the B.A. degree.
http://eeb.yale.edu/sites/default/files/eeb_470_senior_tutorial_form_rev…
1 Yale College course credit(s).
Research (EEB 4474)
One term of original research in an area relevant to ecology or evolutionary biology. This may involve, for example, laboratory work, fieldwork, or mathematical or computer modeling. Students may also work in areas related to environmental biology such as policy, economics, or ethics. The research project may not be a review of relevant literature but must be original. In all cases students must have a faculty sponsor who oversees the research and is responsible for the rigor of the project. Students are expected to spend ten hours per week on their research projects. Using the form available from the office of undergraduate studies or from the Classes server, students must submit a research proposal that has been approved by the faculty sponsor to the director of undergraduate studies, preferably during the term preceding the research. Proposals are due no later than the first day of the second week of the term in which the student enrolls in the course. The final research paper is due in the hands of the of the director of of undergraduate studies by the last day of reading period in the term of enrollment.
Requires Permission of the Instructor
http://eeb.yale.edu/sites/default/files/eeb_474_research_form_rev_8.20.1…
1 Yale College course credit(s).
Senior Research (EEB 4475a/4476b)
One term of original research in an area relevant to ecology or evolutionary biology. This may involve, for example, laboratory work, fieldwork, or mathematical or computer modeling. Students may also work in areas related to environmental biology such as policy, economics, or ethics. The research project may not be a review of relevant literature but must be original. In all cases students must have a faculty sponsor who oversees the research and is responsible for the rigor of the project. Students are expected to spend ten hours per week on their research projects. Using the form available from the office of undergraduate studies or from the Classes server, students must submit a research proposal that has been approved by the faculty sponsor to the director of undergraduate studies, preferably during the term preceding the research. Proposals are due no later than the first day of the second week of the term in which the student enrolls in the course. The final research paper is due in the hands of the director of undergraduate studies by the last day of reading period in the term of enrollment.
Requires Permission from Instructor
Enrollment limited to EEB seniors. Two terms of senior research fulfills the senior requirement for the B.S. degree.
http://eeb.yale.edu/sites/default/files/eeb_475_a_and_b_senior_research_…
1 Yale College course credit(s).
Intensive Senior Research (EEB 4495a/4496b)
One term of intensive original research during the senior year under the sponsorship of a Yale faculty member. Similar to other research courses except that a more substantial portion of a student?s time and effort should be spent on the research project (a minimum average of twenty hours per week). A research proposal approved by the sponsoring faculty member must be submitted to the director of undergraduate studies; forms are available from the office of undergraduate studies. For research in the fall term, approval is encouraged during the spring term of the junior year. Proposals are due no later than the first day of the second week of the term in which the student enrolls in the course. The final research paper is due in the hands of the director of undergraduate studies by the last day of reading period in the term of enrollment.
Requires Permission from Instructor
Enrollment limited to EEB seniors. Two terms of intensive research fulfills the senior requirement for the B.S. degree.
http://eeb.yale.edu/sites/default/files/eeb_495a_eeb_496b_intensive_sr_r…
Advanced Topics in Ecology & Evolutionary Biology (EEB 6500)
Topics to be announced. Graded Satisfactory/Unsatisfactory.
Topics to be announced. Graded Satisfactory/Unsatisfactory.
Responsible Conduct of Research (EEB 6545)
Graded Satisfactory/Unsatisfactory.
Requires Permission of the Instructor
This 5-week discussion seminar considers issues related to the responsible conduct of research. Topics addressed include: research misconduct, plagiarism, data acquisition and management, mentoring and collaboration, authorship and peer review, the use of animals and humans in scientific research, sexual harassment, diversity, and balancing professional and personal life.
This course takes place within EEB 6501: Advanced Topics. First-year students are required to register for both courses.
Research Rotation I (EEB 6901)
Research Rotation
Research Rotation II (EEB 6902)
Research Rotation II
Biology of Insect Disease Vectors (EEB 7650)
Insects transmit pathogens that cause many emerging and re-emerging human and agriculture-related diseases. Many of these diseases, which are referred to as neglected tropical diseases (NTDs), have a dramatically negative impact on human health in the developing world. Furthermore, they cause indirect devastation by significantly reducing agricultural productivity and nutrient availability, exacerbating poverty and deepening disparities. This course introduces students to the biological interactions that occur between major groups of important disease vectors and the pathogens they transmit. Lectures cover current research trends that relate to the ecology and physiology of insect vectors. Course content focuses on how these aspects of vector biology relate to the development and implementation of innovative and effective disease-control strategies.
Prerequisite: full year of college/university-level biology, or permission of the instructor(s).
Foundations of Ecology (EEB 7712)
This seminar course familiarizes students with foundational concepts and themes in ecology and how they have changed over time. Each week we read and discuss two papers: one classic paper selected from the recently published volume Foundations of Ecology II: Classic Papers with Commentaries (Eds. Miller and Travis, 2022) covering the period 1970–1995, and one related contemporary paper published after 2010. We discuss how the concepts and themes introduced in classic papers have influenced the field of ecology and consider how new tools, data, and insights have advanced, diminished, or changed their impact. The Foundations book covers many topics, arranged into six core areas. Readings cover all six areas, but the included content varies depending on the interests of the class. Students are responsible for choosing one classic paper from Foundations, pairing it with one contemporary paper and leading the discussion during the class meeting. Students also submit short weekly “reflections” in response to a prompt.
From Biodiversity Science to Conservation Impact (EEB 7713)
n December 2022, nations worldwide committed to a new set of goals for biodiversity conservation. In the 15th Conference of the Parties of the Convention on Biological Diversity they agreed to a Global Biodiversity Framework that lays out ambitious goals and targets. Importantly, for the first time, the framework specifies biodiversity monitoring to support achieving these goals, setting up a more critical role than ever for relevant science to contribute to the success of the overall ambition. This course addresses the science for measuring progress and providing decision-support for meeting these commitments. Following an initial assessment of framework language, we evaluate for each of the goals and targets that address biodiversity threats the required evidence to support a robust and representative measurement at global scale. We discuss key scientific papers addressing the concepts, methods, and evidence supporting these measurements for ecosystems and species. We then critically evaluate how currently available data, methods, and tools can support the monitoring envisioned by the framework. We closely review the opportunities enabled by remote sensing and the new generation of model-supported indicators of biodiversity change they support. Finally, we discuss the potential of several new or emerging technologies and approaches such as camera traps, GPS tracking, community science, machine learning, and others to offer an improved foundation for monitoring planetary biodiversity change.
Structuralism and Macroevolution (EEB 7717 )
A seminar course discussing the philosophical roots of and empirical research in structuralism and macroevolution. We read selected papers in philosophy of evolutionary biology, comparative phylogenetic methods, macroevolutionary studies, and the role of natural history in evolutionary thought. Each topic is paired with readings on empirical research that involves similar issues. The course concludes with a short writing assignment that analyzes a contemporary question in macroevolution or structural/organismic research.
Foundations of Terrestrial Ecology (EEB 7721)
Intended for graduate students, this seminar course brings a historical perspective to understanding current questions and approaches in terrestrial ecology, ranging from evolutionary, community, landscape, to ecosystem ecology. We read and discuss foundational papers and related current papers, and we identify future directions, opportunities, and challenges for the different sub-fields. The course allows students to critically examine and engage with some scientific work that has laid the findings and concepts that are foundational as they develop conceptual and methodological approaches to their own research. Starting in weeks three or four, each student takes a turn leading discussion and selecting a relevant current paper to that week’s topic. Students write a total of seven précis on the topics of their choosing, with at least two completed by week six.
Comparative Genomics (EEB 7723)
The field of evolutionary biology is increasingly drawing on genomic data, and the field of genomic biology is becoming more evolutionary as genomes are sequenced for a broader diversity of organisms. This course focuses on the evolution of genome sequence and function at macroevolutionary timescales, with an emphasis on building practical computational skills for genomic and phylogenetic comparative analyses. The focus is more on using phylogenies to understand genome evolution than on using genomes to build phylogenies.
Scientific Writing for Ecology and Evolutionary Biology (EEB 7725)
This course will provide guidance and practice for graduate students in grant and manuscript writing in the fields of ecology and evolutionary biology. Students will produce one grant application (NSF GRFP/DDIG or similar) and one manuscript for publication (on a topic of their choice, to contribute to their thesis or other ongoing work). Application to course & instructor permission required: see Canvas site.
Forgotten Grassy Ecosystems (EEB 7750)
Grassy ecosystems—including savannas and grasslands—have historically been relatively undervalued, often confused and misclassified as forests. This seminar includes weekly readings and discussion about the world’s grassy ecosystems in general and focusing on regional examples of overlooked savannas and grasslands. This seminar is intended for Ph.D. students. It is open to master’s students and undergraduates by permission of the instructor only, based on a one- or two-paragraph description of interest in the course.
Ecology of Landforms (EEB 7762)
This course is a combined graduate research seminar and research practicum that explores the linkage between ecological and geomorphic processes—between biology at Earth’s surface and the shape and structure of that surface. This course is centered around two skill-building activities: (1) a series of presentations in which students deliver short mock lectures, lead subsequent discussion, and receive constructive feedback from the class; (2) a series of quantitative workshops (held in alternate weeks) in which the class collaboratively designs and codes a model or model(s) that relate to ecological and geomorphic processes. A primary focus of this course is understanding how biogeochemical cycles play out across hillslopes, watersheds, and fluvial landforms. Depending on student interest we may also address questions relating community ecology, population ecology, or evolutionary processes to landscape structure. Students should expect to hone their presentation skills and quantitative toolset, particularly with respect to spatial analysis and numerical modeling.
Life in the Anthropocene (EEB 7804)
All living things exist in an era of unprecedented global-scale environmental change. Global change encompasses numerous, often interconnected phenomena that are currently impacting organisms. These include rising temperatures, ocean acidification, habitat loss and degradation, overexploitation, novel pathogens, and toxin exposure. This course focuses on global change from the perspective of the organisms themselves. Our goal as biologists is to understand the magnitude of the problem by addressing the following questions: (1) What are the principal ways in which organisms are being challenged due to human impact? (2) What mechanisms are there for organisms to adaptively respond to these challenges? and (3) What can we do to help organisms? To address these questions, we delve into the scientific literature on distinct topics related to global change, discussing one of these topics in each class meeting. Key papers from the literature are assigned to help guide discussion. This course is discussion-based and interactive; each week a student leads discussion along with the instructor. A secondary goal of the course is to help students improve their written communication abilities, either through a traditional term paper or something else, like a grant application or a dissertation chapter. Depending on the number of students and their goals, we might tackle a review paper together, with the goal of obtaining a peer-reviewed publication.
Prerequisites: introductory biology and evolution.
The Ecology of the Great Pandemics (EEB 7830)
In this course we examine principles of the ecology of infectious disease in light of three pandemics: the 1918 influenza pandemic, the HIV/AIDS pandemic, and the COVID-19 pandemic. The course covers principles of zooneses, disease emergence, herd immunity, basic vaccinology, and other fundamental concepts. It also focuses on social and cultural factors that fomented these pandemics.
Adaptive Radiation (EEB 7834)
One of the most striking patterns of biodiversity is its uneven distribution across the tree of life. Theory suggests that this disparity in diversity reflects feedback between ecological and phenotypic evolution. Adaptive radiations, the rapid multiplication of species into distinct ecological niches, is a key example of this phenomenon. While studies about adaptive radiation have grown exponentially over the past decades, fundamental questions remain. For example, how does phenotypic diversification unfold during radiation? How do species interactions shape species richness in adaptive radiation? Do adaptive radiations play out in parallel across continental and island contexts, or are there repeatable differences among them? The goal of this course is to critically dissect the field of adaptive radiation. Specifically, we unpack its major features and identify unresolved lacunae. To do so, we delve into the scientific literature on distinct topics related to adaptive radiation. Key papers from the literature are assigned to help guide discussion. We begin with a brief survey of recent syntheses on adaptive radiation to refresh our understanding and, most importantly, identify key gaps in empirical knowledge. Then, we explore several studies focused on less-studied axes of adaptive radiation to catalyze our thinking.
Prerequisites: BIOL 103/104 (or equivalent) and EEB 5225 (or equivalent).
The Behavioral Immune System (EEB 7854)
Behavior is the first line of defense against parasites and pathogens. Behavioral defenses allow organisms to minimize contact with infectious agents, and the concept of the “behavioral immune system” was developed to encompass a range of evolved behaviors that help minimize the fitness costs of infection. The COVID-19 pandemic has made the term “social distancing” a household term; however, distancing and many other avoidance strategies are employed by a wide range of organisms to combat infectious agents. In this seminar, we examine our current understanding of the behavioral immune system across the diversity of animals, including humans. Specifically, we explore: (1) the mechanisms of behavioral immunity; (2) the ecological, evolutionary, and epidemiological consequences of these behaviors; and (3) key costs of behavioral immunity that maintain intra- and interspecific variation. To do this, we discuss and synthesize the scientific literature on the behavioral immune system, drawing parallels to work on the physiological immune system. The first weeks of the course focus on instructor-selected papers, and subsequent weeks incorporate student-selected papers.
Special Topics in the Ecology and Evolution of Infectious Diseases (EEB 7856)
Historically, pathogens and the diseases they cause were viewed largely from a biomedical perspective focused on interactions between pathogens and their human hosts. However, in the last few decades, the importance of studying pathogens from an ecological and evolutionary perspective has gained significant traction. These perspectives inform our understanding of almost all aspects of pathogen-host interactions from transmission dynamics and zoonotic disease spillover to the evolution of virulence and drug resistance. In this seminar, we dissect current and classic literature on the ecology and evolution of infectious diseases. Specifically, we: (i) discuss fundamental concepts in the field; (ii) identify persistent knowledge gaps; and (iii) explore opportunities for linkages between ecological, evolutionary, and biomedical perspectives.
Ecosystem Dynamics of Nature-Based Climate Solutions (EEB 7862)
Nature-based climate solutions have gained increasing attention in the past decade as possible contributors to reducing our net carbon emissions to the atmosphere, without necessarily reducing gross emissions. Prominent nature-based solutions include forestation (reforestation, afforestation, plantation forestry), avoided deforestation, and soil carbon sequestration. This seminar includes weekly readings and discussion around themes in the management of ecosystem carbon storage.
This seminar is intended for Ph.D. students. It is open to master’s students and undergraduates by permission of the instructor only, based on a one- or two-paragraph description of interest in the course.
Evolutionary Architects: Organisms as Targets and Agents of Natural Selection (EEB 7865)
Organisms are routinely faced with many abiotic and biotic pressures that impact their survivorship, growth, and reproductive success. For example, a lizard’s ability to perform fitness-based tasks (like foraging or predator evasion) is limited by the thermal dependence of its performance, its hydric and metabolic economy, and its morphological dimensions. Yet, organisms are not exclusively at the whim and mercy of their surroundings. Of key importance is the preeminent role that organisms exert on their own selective environments and, correspondingly, on their evolution. This course considers the diverse ways in which organisms engineer their own evolutionary trajectories. Some of the topics we cover include niche construction, extended phenotypes, behavioral drive, the Bogert effect, and adaptive virulence (particularly in the context of the COVID-19 pandemic).
Open to upper-level undergraduates who have taken BIOL 1030 (link is external), BIOL 1040 (link is external), and EEB 2225 (link is external) (or the equivalent).
Ecological Genomics (EEB 7870)
Speciation & Adaptation Genomics (EEB 7872)
Speciation and adaptation are two fundamental processes that generate the diversity of life seen on earth to date. This graduate-level seminar course will explore the evolutionary mechanisms responsible for these phenomena by delving into the primary literature to explore classic examples of adaptation and speciation using a genetics and genomics lens.
Seminar in Systematics (EEB 7930)
Topics and class time are chosen by the participants, and have included reading books and/or a series of papers on particular topics (e.g., homology; morphological phylogenetics; evolution of egg colors and exposed nesting in dinosaurs/birds; origin of snake ecology; conflicts between morphology and molecules; role of fossils in phylogenetic inference).