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Studies problems in demography, epidemiology, ecology, and social behavior of human and nonhuman populations, using tools of mathematics, computation, and statistics. In demography, he studied the uncertainty of population projections theoretically and in practical applications ranging from striped bass in the Chesapeake Bay to asbestos-related diseases in humans. His analysis of how many people the Earth can support found that population, economics, the environment, and culture jointly influence trade-offs in human well-being. In epidemiology, he developed mathematical models of malaria, schistosomiasis, and Chagas disease. In ecology, he proposed and tested new models of food webs.