Joan Roughgarden is an American ecologist, evolutionary biologist, and author. Roughgarden is also one of the most notable transgender scientists in history.
Some of Roughgarden’s work has included scientific criticism of scholarship that affects sex and gender minorities. This profile focuses on that work, particularly Roughgarden’s important role in the community response to the anti-transgender book The Man Who Would Be Queen by J. Michael Bailey, published in 2003.
Background
Joan Elizabeth Roughgarden was born on March 13, 1946 in Paterson, New Jersey. Roughgarden earned bachelor’s degrees from University of Rochester in 1968 and a doctorate from Harvard University in 1971. Roughgarden joined Stanford University in 1972, soon founding and directing the Earth Systems Program. Roughgarden was named a Fellow of the Gugenheim Foundation in 1985 and elected a Fellow of the American Academy of Arts and Sciences in 1993. Roughgarden made a gender transition in 1998, coming out at age 52.
Roughgarden was named a Fellow at Stanford in 1978 and became professor emerita at Stanford in 2011. Roughgarden is also affiliated with the Hawaiʻi Institute of Marine Biology.
Work and views
Roughgarden argues that traditional sexual selection theory overemphasizes male competition and female choice, projecting overly Darwinian conflict models onto animal behavior. Roughgarden contends that many species instead exhibit cooperation, social negotiation, and mutual mate choice, which the classic framework fails to explain. Roughgarden proposes “social selection” as a broader alternative that accounts for cooperative reproductive strategies and diverse sexual systems.
Roughgarden disputes the gene-centered view of evolution promoted by Richard Dawkins and others, arguing that it treats organisms as passive vehicles for “selfish” genes and oversimplifies how traits evolve. Roughgarden contends that cooperation and social negotiation play a larger role in evolution than Dawkins’s framework allows and has suggested the alternative “genial” gene. Their disagreement reflects a deeper clash over whether evolutionary theory should prioritize competition or incorporate broader models of cooperation and social complexity.
Roughgarden’s work is often criticized by people whom Stephen Jay Gould called “Darwinian fundamentalists,” especially anti-trans biologists like Dawkins, anti-trans linguists like Steven Pinker, and evolutionary psychologists like J. Michael Bailey.
Roughgarden is also a proponent of theistic evolution, described in the 2006 book Evolution and Christian Faith.
The Man Who Would Be Queen (2003)
Roughgarden was a key critic of author J. Michael Bailey’s 2003 book The Man Who Would Be Queen. Roughgarden’s first-hand report of Bailey’s vulgar misuse of gender diverse children while on book tour at Stanford was a galvanizing moment for trans people worldwide. Calling it “the most humiliating lecture I’ve ever personally attended,” in a 2003 letter, Roughgarden called for the National Academies (NASEM) to recall the book and release it for publication elsewhere.
In 2003, J. Michael Bailey published a response on a Northwestern University faculty page:
Joan Roughgarden, who is a transsexual biology professor at Stanford University, attempted to get me uninvited from giving a talk to the Stanford Psychology Department. (So much for open scientific debate.) When that attempt failed, she attended my talk and wrote an insulting and scathing review for the Stanford paper. The talk was not about transsexualism, per se, but about sexual orientation and gender nonconformity. To say that I disagree with Roughgarden’s take on the lecture is an understatement, but I only want to address two points.
First, regarding the tone of the lecture, and my attitude toward gender nonconforming homosexual people: I invite anyone who is concerned to read my book, The Man Who Would Be Queen, particularly the first two sections, which are most relevant. Any reasonable person will conclude that I am very sympathetic to the plight of gender nonconforming boys, and that my research is intended to illuminate gender nonconformity as a truly fascinating phenomenon. When some in the Stanford audience giggled at some of the demonstrations in my talk (e.g., my playing the voices of gay and straight people), this was all in good humor. A gay psychologist and sex researcher, James Cantor, wrote in response to Roughgarden’s screed:
“I have seen Bailey give this lecture before (at least, an earlier version of it). Again, this was the one with several openly lesbian women and gay men in the audience, including me. None of us felt at all offended. What Roughgarden describes as laughter was actually an affectionate recognition of the truth. Effeminate speech is much more common among gay men than straight men, and telling the two extremes apart is like night and day.”
Second, it would not surprise me if some gay or lesbian people in the audience were uncomfortable with the talk. I have written about the discomfort that some gay men, particularly, feel about linking male homosexuality and femininity (and this link is a major idea in my book). I discuss gay men’s discomfort with the idea of gay femininity in my book. The fact is, however, that this link is empirically well established and scientifically interesting. Furthermore, there is nothing wrong with feminine gay men. It is the discomfort that some gay men feel with this association that is the problem, not my ideas. And I would like everyone to become more comfortable with male femininity.
A presentation and book signing at A Different Light bookstore in Los Angeles was cancelled “at the request of many in the local transgendered community.” I am certain that more incidents such as this will occur, and that more criticism like Roughgarden’s and Conway’s will be offered. So be it.
The controversy has already consumed substantial time that I could be spending on new research, teaching, and administration, and I cannot afford more time to respond to each new charge made by Conway, Roughgarden, et al. Please feel free to send an email to me, however. I am most likely to respond to comments that suggest a serious effort to understand my arguments (especially by reading my book).
In a 2004 review in Times Higher Education, Roughgarden found the book racist, homophobic, and misogynist:
Bailey’s writing seems to me hate speech, detached from reality, and yet advertised as science and published by the US National Academies.
Bailey has manipulated even the few narratives he has. He admits to an “ongoing argument” with one of his subjects who will not agree to say what he wants. When his subjects disagree with him he implies they are liars. “Most gender patients lie”, according to an “ace gender clinician” Bailey quotes. Also, gay males who do not report a feminine childhood are lying too because they suffer from “internalised femiphobia”. Therefore, Bailey corrupts what data he has by putting words in the mouths of his interviewees.
As of August 2003, his six transgendered subjects have each filed formal complaints at Northwestern charging that Bailey did not notify them that their narratives were to be used as “research material” in his book (“Transsexuals file 2 more claims against Bailey”, Daily Northwestern, August 2 2003).
After reading accounts supposedly about them, the women reported being inaccurately transcribed. In December 2003, one of the women formally charged Bailey with having sex with her at her apartment (“Northwestern U. psychologist is accused of having sex with research subject”, Chronicle of Higher Education, December 12 2003.) Bailey has declined to comment. He also did not disclose that he was writing letters for these women to authorise sex-reassignment surgeries in return for their interviews, placing the women and him in conflict of interest. And in April this year, a formal complaint was filed by three prominent transgender activists with the Illinois Department of Professional Regulation and with Northwestern charging that Bailey was functioning as a clinical psychologist without a licence. At the time of writing this review, none of these complaints has been resolved.
Last, but not least, the major narrative that frames the section in the book on transgendered women (the “Danny” narrative) is acknowledged to be fabricated. Thus, all of Bailey’s narrative data are irretrievably compromised and the practices with which the information was obtained are allegedly scandalous.
All in all, Bailey’s book is not only politically incorrect, it is scientifically incorrect.
Roughgarden appeared on KQED for a 2007 Forum episode with host Michael Krasny called Transgender Theories. That episode included Mara Keisling, Bailey, and Alice Dreger, who had just gotten Benedict Carey of the New York Times to promote Dreger’s cover-up of Bailey’s academic misconduct. Stanford neurobiologist Ben Barres calls in during the episode.
Evolution’s Rainbow (2004)
Roughgarden laid out the latest research on sex and gender diversity in the 2004 book Evolution’s Rainbow: Diversity, Gender, and Sexuality in Nature and People. Roughgarden commented on “scientific” descriptions of reptile behavior that were not value-neutral. Roughgarden showed how bias about transgender people crept into descriptions of garter snakes and singled out use of the term “she-male” as an especially biased example.
These studies are disturbing because they attempt to sensationalize at the expense of transgendered people. An article in the scientific journal Animal Behaviour begins, “Female mimicry, whereby a male takes on a female’s appearance, is a rare but widely publicized trait in human societies. Remarkably, parallels can be seen in other animal species.” Feminine males and masculine females are not rare among humans. Nor are transgendered people comparable to snakes. All male garter snakes wear female perfume and participate in same-sex copulation every year. No human society has ever enjoyed such a rite of spring! An article in another scientific journal. the Canadian Journal of Zoology, refers to female mimicry as “bizarre.” The problem with female mimicry is not that it is bizarre; the problem is that female mimicry is a myth.
Both articles refer to feminine males in text and figure captions as “she-males.” This language, derived from pronography, is derogatory. A she-male is a woman with a penis. The transgender community has better words to describe transgendered bodies. In the biological literature expressions like “gynomorphic male” and “andromorphic female” are preferred when describing a feminine male or masculine female.
The articles derogate not only transgendered people but also their partners. The title of one article includes the phrase “transvestite serpent,” and another claims to be about the “behavioural tactics of ‘shemales’ and the males who court them.” This writing not only stigmatizes transgendered people with certain body types, but also transfers the stigma to their friends. I hope future work on these animals is carried out with more professionalism, and that future publications on this subject receive better editorial oversight.
ProSocial World article series (2025)
In 2025, Roughgarden introduced a planned series of articles about biology, sex, and gender for ProSocial World, a nonprofit founded in 2020 to combine scientific theory with practical tools. On release of the project, Roughgarden and Jaimie Veale also published an opinion piece outlining their views.
References
MIT Press (February 11, 2019). Q&A with Joan Roughgarden on the Problems with the Theory of Sexual Selection. MIT Press https://mitpress.mit.edu/blog/qa-joan-roughgarden-problems-theory-sexual-selection [archive]
Surkan KJ (2007). Transsexuals Protest Academic Exploitation.
Dorit, Robert (September 2004). Rethinking Sex. American Scientist https://www.americanscientist.org/article/rethinking-sex
Moser, Bob (May 2004). On the Originality of Species: Biologist Joan Roughgarden has studied nature and thinks the rule needs to change. That means challenging Darwin on sex. Stanford Magazine https://stanfordmag.org/contents/on-the-originality-of-species
Bailey, J. Michael (2003). Book Controversy Question & Answer. http://faculty.wcas.northwestern.edu/JMichael-Bailey/controversy.htm
Yoon, Carol Kaesuk (October 17, 2000). Scientist at Work: Joan Roughgarden; A Theorist With Personal Experience Of the Divide Between the Sexes. New York Times https://www.nytimes.com/2000/10/17/science/scientist-work-joan-roughgarden-theorist-with-personal-experience-divide-between.html
Selected writing by Roughgarden
Roughgarden, Joan (November 11, 2025). Sex, Gender Diversity, and Evolution: Introduction ProSocial World https://www.prosocial.world/posts/sex-gender-diversity-and-evolution-introduction
Roughgarden, Joan; Veale, Jaimie (November 19, 2025). We’re scientists who study sex and gender. Here’s what Trump gets wrong about trans people. San Francisco Chronicle https://www.sfchronicle.com/opinion/openforum/article/trans-gender-sex-female-male-21125145.php alt headline: What President Trump gets wrong about transgenderism proposed headline: Transgender is not ideology.
Roughgarden, J. (2017). Homosexuality and Evolution. In On Human Nature (pp. 495–516). Elsevier. https://doi.org/10.1016/b978-0-12-420190-3.00030-2
Roughgarden, J. (2015). Sexual Selection: Is Anything Left?. In: Hoquet, T. (eds) Current Perspectives on Sexual Selection. History, Philosophy and Theory of the Life Sciences, vol 9. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-9585-2_5
Roughgarden, J. (2012). The social selection alternative to sexual selection. Philosophical Transactions of the Royal Society B: Biological Sciences, 367(1600), 2294–2303. https://doi.org/10.1098/rstb.2011.0282
Roughgarden, J., & Akçay, E. (2010). Do we need a Sexual Selection 2.0? Animal Behaviour, 79(3), e1–e4. https://doi.org/10.1016/j.anbehav.2009.06.006
Milam, E. L., Millstein, R. L., Potochnik, A., & Roughgarden, J. E. (2010). Sex and sensibility: The role of social selection. Metascience, 20(2), 253–277. https://doi.org/10.1007/s11016-010-9464-6
Roughgarden, Joan (2007). Challenging Darwin’s theory of sexual selection. Daedalus Volume 136 | Issue 2 | Spring 2007 p.23-36 https://doi.org/10.1162/daed.2007.136.2.23
oughgarden, J., Oishi, M., & Akçay, E. (2006). Reproductive Social Behavior: Cooperative Games to Replace Sexual Selection. Science, 311(5763), 965–969. https://doi.org/10.1126/science.1110105
Roughgarden, J. (2004). Evolution and the Embodiment of Gender. GLQ: A Journal of Lesbian and Gay Studies, 10(2), 287–291. https://doi.org/10.1215/10642684-10-2-287
Roughgarden, Joan (May 28, 2004). Lots of sex but a lack of science: The Man Who Would Be Queen, Times Higher Education. https://www.timeshighereducation.com/books/lots-of-sex-but-a-lack-of-science/189033.article
Roughgarden, Joan (February 11, 2004). The Bailey Affair: Psychology Perverted. https://www.researchgate.net/profile/Joan-Roughgarden/publication/266863785_The_Bailey_Affair_Psychology_Perverted/links/546411bd0cf2cb7e9da99e22/The-Bailey-Affair-Psychology-Perverted.pdf
Books
Roughgarden J (2009). The Genial Gene: Deconstructing Darwinian Selfishness. University of California Press ISBN 978-0-520-25826-6
Roughgarden J (2006). Evolution and Christian Faith: Reflections of an Evolutionary Biologist. Island Press ISBN 1-59726-098-3
Roughgarden J (2004). Evolution’s Rainbow: Diversity, Gender and Sexuality in Nature and People. University of California Press ISBN 0-520-24073-1
Roughgarden J (1997). Primer of Ecological Theory. Prentice Hall
Roughgarden J (1995). Anolis Lizards of the Caribbean: Ecology, Evolution and Plate Tectonics. Oxford University Press
Roughgarden J, May RM, Levin SA, Eds. (1995). Perspectives in Ecological Theory. Oxford Univ. Press
Ehrlich PR, Roughgarden J (1987). Science of Ecology. Prentice Hall
Roughgarden J (1979). Theory of Population Genetics and Evolutionary Ecology: an Introduction. Prentice Hall
Media
Joan Roughgarden (May 19, 2025). Deep Dive Contrasting Social and Sexual Selection – Roughgarden. https://www.youtube.com/watch?v=yBqalBf4_GM
Joan Roughgarden (April 30, 2025). Trans People Across Cultures and Through History –Joan Roughgarden 04/08/2025, 52 min with Q&A. https://www.youtube.com/watch?v=GCyJskyEyIE
Joan Roughgarden (June 13, 2022). Variation in Gender Expression and Sexuality: Implications for Evolutionary Theory. https://www.youtube.com/watch?v=ypQKaYaM7BY
Outwords with Joan Roughgarden (August 30, 2021). Joan Roughgarden. https://theoutwordsarchive.org/interview/joan-roughgarden/
Joan Roughgarden (March 11, 2016). Roughgarden on the Gender Binary in Nature, Culture and the Bible. https://www.youtube.com/watch?v=8sdjnheBMCI
Joan Roughgarden (March 12, 2014). Roughgarden on Jesus & Darwin, 1st Malcolm Miner Lecture, St. Michaels, Kauai HI. https://www.youtube.com/watch?v=peH2yAyfcOE
Joan Roughgarden (May 22, 2012). Roughgarden on Evolution of Gender and Sexuality. https://www.youtube.com/watch?v=x80WXeP-2vI
Joan Roughgarden (May 6, 2012). Roughgarden on Evolution and Christianity. https://www.youtube.com/watch?v=AtJ2KfZQfLk
Joan Roughgarden (May 4, 2012). Roughgarden on Social Selection. https://www.youtube.com/watch?v=sf-Ya-X1iLE
Joan Roughgarden (March 27, 2012). Roughgarden on Sexual and Social Selection. https://www.youtube.com/watch?v=i_V8njOdHlI
TEDx Talks (February 24, 2011). Sexual diversity in nature: Joan Roughgarden at TEDxAmazonia. https://www.youtube.com/watch?v=KJ3lcCa0G4Y
Barry Belmont (Jan 11, 2011). 18. Joan Roughgarden (1 of 2) – Beyond Belief 2006. https://www.youtube.com/watch?v=-gbFLml1uvM
- 19. Joan Roughgarden (2 of 2) – Beyond Belief 2006. https://www.youtube.com/watch?v=6dNRAlkieIo
- 20. Richard Dawkins – Beyond Belief 2006. https://www.youtube.com/watch?v=iijmYn1KbFs
- 21. Joan Roughgarden and Richard Dawkins Take Questions (1 of 2) – Beyond Belief 2006. https://www.youtube.com/watch?v=8EY7JYAIW6s
- 22. Joan Roughgarden and Richard Dawkins Take Questions (2 of 2) – Beyond Belief 2006. https://www.youtube.com/watch?v=lzoUudp5Auk
- 23. Roughgarden, Dawkins, Harris, and Tyson Discussion (1 of 2) – Beyond Belief 2006. https://www.youtube.com/watch?v=yeYdLe6A2bc
- 24. Roughgarden, Dawkins, Harris, and Tyson Discussion (2 of 2) – Beyond Belief 2006. https://www.youtube.com/watch?v=rfOuK2fKjqI
GenderTalk with Nancy Nangeroni and Gordene MacKenzie (June 16, 2003). Program #Â 414: Joan Roughgarden. https://www.gendertalk.com/radio/programs/400/gt414.shtml
Resources
Wikipedia (en.wikipedia.org)
Stanford University (stanford.edu)
- Roughgarden Lab [archive]
- web.stanford.edu/group/roughlab
- Joan Roughgarden [archive]
- web.stanford.edu/~rough
- Joan Roughgarden CV (as of July 15, 2020)
- web.stanford.edu/~rough/Roughgarden-CV-20200715.html
- Joan Roughgarden
- profiles.stanford.edu/joan-roughgarden
joandistrict6.com [archive]
Ram-2050 (ram-2050.com) [archive]
Facebook (facebook.com)
LinkedIn (linkedin.com)
X/Twitter (x.com)
YouTube (youtube.com)
Note: In 2025, this site phased out AI illustrations after artist feedback. The previous illustration is here.