Tracy Heath

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Assistant Professor
Dept. Ecology, Evolution, & Organismal Biology
Iowa State University

Contact Information



I develop statistical models for use as priors in Bayesian methods for phylogenetic inference. In particular, I'm interested in divergence-time estimation -- e.g. modeling rate variation across the tree of life, diversification processes, and methods for incorporating information from the fossil record.

Time at MBL

Monday July 24 - Sunday July 30

2017 Course

Morning lecture slides and example scripts

  • Hierarchical models and integrative Bayesian analysis (slides)
    • RevBayes archery model demo (files)
    • RevBayes GTR demo (files)
  • Bayesian divergence-time estimation (slides)

Afternoon tutorial

Suggested Reading

Graphical Models

  • Höhna, Heath, Boussau, Landis, Ronquist, Huelsenbeck. 2014. Probabilistic graphical model representation in phylogenetics. Systematic Biology, 63:753–771. (doi: 10.1093/sysbio/syu039).

Divergence-Time Estimation

  • Thorne and Kishino. 2005. Estimation of divergence times from molecular sequence data. In: Nielsen R, editor, Statistical Methods in Molecular Evolution. New York: Springer, pp. 235–256. (PDF)
  • Heath and Moore. 2014. Bayesian inference of species divergence times. In: Bayesian Phylogenetics: Methods, Algorithms, and Applications. Chen, M.-H., L. Kuo, and P.O. Lewis (editors). Chapman & Hall/CRC. (book chapter email me for PDF)
  • BEAST2 Book: Drummond AJ, Bouckaert RR. 2014. Bayesian evolutionary analysis with BEAST 2. Cambridge University Press. (
  • Lartillot: blog post on diversification:
  • also see references in divergence-time tutorial

Additional Tutorials

  • More tutorials on divergence time estimation in RevBayes and BEAST2 (link)
  • Materials (tutorials and lectures) from the Taming the BEAST workshop, including some tutorials on infectious disease dynamics (link)

2016 Course