Joan Roughgarden

The Gender Binary in Nature and Across Human Cultures

The animal kingdom offers no support for a natural universal distinction between male and female sexual categories for whole organisms. Nor does the animal kingdom support the universal existence of two fixed genders within species. The discovery of extensive variation in gender expression and sexuality in nature challenges traditional biological explanations of animal behavior, especially accounts that trace to Darwin’s theory of sexual selection. Variation in human gender expression and sexual orientation occurs among people in all cultures throughout the world. All cultures have indigenous institutions and norms to accommodate the natural variation in the human species. In Western culture too, even the Bible contains explicit inclusionary reference to human variation in gender expression.

Research Areas

Evolution of cooperation


Joan Roughgarden received bachelor degrees in biology and philosophy from University of Rochester in 1968 and a Ph.D. in biology from Harvard University in 1971. She has taught at Stanford University since 1972 as Professor of Biology and Geophysics. In 2011 she retired and moved to the island of Kauai in Hawaii where she remains a Stanford professor (emerita). In 2012 she joined the University of Hawaii’s Institute of Marine Biology as Adjunct Professor and Research Scientist. While at Stanford, in 1992 Joan founded and directed the Earth Systems Program and in 1995 received the Dinkelspiel Award for service to undergraduate education.


Roughgarden, Joan. 2015. „Sexual selection: Is anything left?“. In: Hoquet, Thierry. Ed. „Current Perspectives on Sexual Selection. What’s Left After Darwin?“. Series: History, Philosophy and Theory of the Life Sciences 9. Dordrecht: Springer Verlag. 85–102.
Roughgarden, Joan. 2014. „Evolution’s Rainbow: Diversity, Gender, and Sexuality in Nature and People“. Tenth Anniversary Edition. Berkeley CA: University of California Press.
Roughgarden, Joan. 2009. „The Genial Gene: Deconstructing Darwinian Selfishness“. Berkeley CA: University of California Press. Berkeley CA.


Joan Roughgarden
Hawaii Institute of Marine Biology
University of Hawaii
5241 Wili Road
Kapaa HI 9674
Tel.: +1808-415-867-4786


Roughgarden on the Gender Binary in Nature, Culture and the Bible
RAM – 2050