Sean H. Rice

Dept. of Biological Sciences
Texas Tech University
Lubbock, TX 79409
Phone: (806) 742-2710 ext. 274 (Lab: ext 275)

General academic interests and approaches

My interests reach throughout evolutionary biology and my work has made contact a wide range of topics, including developmental modeling, morphological evolution in both invertebrates and vertebrates, population and quantitative genetics, macroevolution, and the philosophical and conceptual basis of evolutionary theory. My theoretical work currently focuses on two broad areas: 1) developing a rigorous theoretical framework for the study of developmental evolution, and 2) the mathematical and conceptual foundations of evolutionary theory.

This work is united not by a focus on any particular system or method (though I keep coming back to some) but by a guiding philosophy concerning how theories in biology should be constructed. This philosophy emphasizes the importance of choosing the right abstractions and understanding their biological meaning. The "theory" is not the equations per se, but rather the way that we assign causal biological interpretations to the abstract terms that the equations are built out of. In this view, doing theoretical biology goes hand in hand with a development of a deep understanding of the biology of the organisms under study. This philosophy also favors analytical models over computer simulations, since the very structure of an analytical model gives insights into biology.

Empirical research by myself and my students focuses on systems that allow us to address important conceptual issues. Recent and current empirical projects in my lab include: the role of heterochrony in human brain evolution, patterns of evolution in the vertebrate skull, morphological evolution in snails and ammonites, and the mechanics of speciation in rotifers.

More discussion of some recent and ongoing projects is given below.

Click on a figure for a more extensive discussion with pictures:

Current projects

An axiomatic theory of evolution : The basic goal of this project is to start with a small set of basic biological premises (termed 'scientific axioms') and derive from these the exact mathematical rules underlying all evolutionary processes, both deterministic and stochastic.

Modeling evolution with epistasis : I am developing a mathematical framework with which to model the evolution of a phenotypic character through modifications of the developmental processes that construct that character. I am using this to investigate the evolution of canalization, integration of characters, and changes in development that are not manifest in adult phenotype.

Some previous projects

Heterochrony : I have proposed a restricted definition of heterochrony that allows us to specify exactly what sorts of morphological transformations are, or are not, heterochrony, and what this means biologically. Using this concept, and a statistical test based on it, I have been investigating the role of heterochrony in primate brain evolution.

Developmental modeling of Moluscan shells : Using a model based on the processes by which mollusc shells are built, we are studying how development influences patterns of morphological evolution in gastropods and the evolution of irregular forms among ammonites.

Multi-level selection: Selection can act at many levels of organization simultaneously. My work focusses on how this influences evolutionary dynamics as well as on foundational issues in evolutionary theory.

Evolutionary Theory: Mathematical and Conceptual Foundations

2004. Published by Sinauer Associates
You can order this book from

Evolutionary Theory is for graduate students, researchers, and advanced undergraduates who want an understanding of the mathematical and biological reasoning that underlies evolutionary theory. The book covers all of the major theoretical approaches used to study the mechanics of evolution, including classical one- and two-locus models, diffusion theory, coalescent theory, quantitative genetics, and game theory. There are also chapters on theoretical approaches to the evolution of development and on multilevel selection theory. Each subject is illustrated by focusing on those results that have the greatest power to influence the way that we think about how evolution works. These major results are developed in detail, with many accompanying illustrations, showing exactly how they are derived and how the mathematics relates to the biological insights that they yield. In this way, the reader learns something of the actual machinery of different branches of theory while gaining a deeper understanding of the evolutionary process.

Roughly half of the book focuses on gene-based models, the other half being concerned with general phenotype-based theory. Throughout, emphasis is placed on the fundamental relationships between the different branches of theory, illustrating how all of these branches are united by a few basic, universal, principles.

The only mathematical background assumed is basic calculus. More advanced mathematical methods are explained, with the help of an extensive appendix, when they are needed.

Here is a list of typographical errors that I have found in the book. Please let me know if you find any others.


Here is a list of some downloadable papers.

My graduate students devise their own research projects. I take the title "advisor" seriously; I interact extensively in identifying a subject and shaping a practical research program, but dissertation projects undertaken by my students are not offshoots of my own research.

Current Grad Students

John Harting
Multivariate evolutionary theory.

Ben Qin
Genome evolution and complexity.

Sarah Fumagalli
Evolution of cooperation


Former Graduate Students

Lisa Suatoni
Evolution of the mechanisms underlying reproductive isolation in rotifers.

Heinrich Zu Dohna (joint with Forestry)
Plant/Insect interactions.

David Ralph
Cultural evolution

Morgan Baker

Former Postdocs

Paul Magwene
Currently Assistant Professor at Duke University.

Anthony Popadopoulos


My principal courses are:

Some Related Sites

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Page last updated on Jul 8, 2021