Ph.D. 1971, Harvard University
M.A. 1965, Harvard University
B.Sc., A.R.C.S. 1963, Imperial College, University of London


Growth, form and function of animal skeletons, especially shells of bivalved molluscs.

Current projects: pattern of exploitation of potential skeletal designs, paleobiology and evolution of arcoid bivalves, evolution of complexity, significance of the Galapagos in writings of Darwin, Melville and Vonnegut.

Course Information

Historical geology, paleontology (emphasis on quantitative methods), evolutionary biology, and the evolution of complex systems in nature.