A Primer of Evolution

A Primer of Evolution is an open educational resource designed for upper-level college students. It provides a succinct introduction to evolutionary thought revolving around theory, evidence, and practice:

  • It introduces some of the theoretical cornerstones and core concepts of modern evolutionary biology. The goal is for students to be able to apply these concepts and articulate testable hypotheses that explain natural phenomena from an evolutionary perspective.
  • It highlights the diversity of empirical approaches and lines of evidence that scientists use to address evolutionary hypotheses.
  • It helps students to practice approaching problems like scientists and evaluate data to address evolutionary hypotheses. To do so, students learn how to program in R to analyze and visualize data and articulate your interpretations and conclusions.

Each chapter provides a conceptual introduction to the topic and includes R-based exercises that allow students to visualize relevant datasets to practice the testing of evolutionary hypotheses. To help with the R exercises, each chapter also provides additional background on case studies and R programming tutorials that help students to develop the necessary skills.

A Primer of Evolution is entirely written in R Markdown and licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License. Anyone is free to copy and redistribute the material in any medium or format, and to remix, transform, and build upon the material provided they give the original author credit (attribution), they are not using the material for commercial purposes (NonCommerical), and they distribute their work under the same license as the original (ShareAlike). If you would like to adopt and use this resource in any way, feel free to contact Michi Tobler to obtain a copy of all source files.

A Primer of Evolution was made possible with the support of a grant from the The Open/Alternative Textbook Initiative at Kansas State University.


  1. What Evolution Is
  2. Evidence for Evolution
  3. A Mechanism for Change
  4. The Raw Materials for Evolution
  5. Evolutionary Mechanisms I: Modeling Selection
  6. Evolutionary Mechanisms II: Mutation, Genetic Drift, Migration, and Non-Random Mating
  7. Evolution of DNA Sequences
  8. Evolution of Quantitative Traits
  9. Adaptation and Phenotypic Plasticity
  10. Social Behavior and Sexual Selection
  11. Speciation
  12. Evolutionary Medicine I: Aging and Diseases of Civilizations
  13. Evolutionary Medicine II: Evolving Pathogens
  14. Human Evolution