Joan Roughgarden (born Jonathan David Roughgarden, 13 March 1946) is an American
ecologist
Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
and
evolutionary biologist
Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life for ...
. She has engaged in theory and observation of coevolution and competition in ''
Anolis
''Anolis'' is a genus of anoles (), iguanian lizards in the family Dactyloidae, native to the Americas. With more than 425 species, it represents the world's most species-rich amniote tetrapod genus, although many of these have been proposed to ...
'' lizards of the
Caribbean
The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...
, and recruitment limitation in the rocky intertidal zones of California and Oregon. She has more recently become known for her rejection of
sexual selection
Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
and her
theistic evolution
Theistic evolution (also known as theistic evolutionism or God-guided evolution) is a theological view that God creates through laws of nature. Its religious teachings are fully compatible with the findings of modern science, including biological ...
ism.
Personal life and education
Roughgarden was born in
Paterson, New Jersey
Paterson ( ) is the largest City (New Jersey), city in and the county seat of Passaic County, New Jersey, Passaic County, in the U.S. state of New Jersey.University of Rochester
The University of Rochester (U of R, UR, or U of Rochester) is a private research university in Rochester, New York. The university grants undergraduate and graduate degrees, including doctoral and professional degrees.
The University of Roc ...
in 1968 and later a Ph.D. in biology from
Harvard University
Harvard University is a private Ivy League research university in Cambridge, Massachusetts. Founded in 1636 as Harvard College and named for its first benefactor, the Puritan clergyman John Harvard, it is the oldest institution of higher le ...
in 1971. In 1998, Roughgarden came out as
transgender
A transgender (often abbreviated as trans) person is someone whose gender identity or gender expression does not correspond with their sex assigned at birth. Many transgender people experience dysphoria, which they seek to alleviate through tr ...
and changed her name to Joan, making a
coming out
Coming out of the closet, often shortened to coming out, is a metaphor used to describe LGBT people's self-disclosure of their sexual orientation, romantic orientation, or gender identity.
Framed and debated as a privacy issue, coming out of ...
post on her website on her 52nd birthday.
Career
Roughgarden worked as an instructor and Assistant Professor of Biology at the
University of Massachusetts Boston
The University of Massachusetts Boston (stylized as UMass Boston) is a Public university, public research university in Boston, Massachusetts. It is the only public research university in Boston and the third-largest campus in the five-campus Un ...
from 1970 to 1972. In 1972 she joined the faculty of the Department of Biology at
Stanford University
Stanford University, officially Leland Stanford Junior University, is a private research university in Stanford, California. The campus occupies , among the largest in the United States, and enrolls over 17,000 students. Stanford is consider ...
. After becoming full professor she retired in 2011, and became Emeritus Professor. She founded and directed the Earth Systems Program at Stanford and has received awards for service to undergraduate education. In 2012 she moved to Hawaii, where she became an adjunct professor at the
Hawaiʻi Institute of Marine Biology
The Hawaii Institute of Marine Biology (HIMB) is a marine biology laboratory located on the state-owned Coconut Island in Kāneohe Bay.
History
The institute was established in 1912. It began as a wooden structure on the shores of Waikiki and o ...
. In her academic career, Roughgarden advised 20 Ph.D. students and 15 postdoctoral fellows.
Roughgarden has authored books and over 180 scientific articles. In addition to a textbook on ecological and evolutionary theory in 1979, Roughgarden has carried out ecological field studies with Caribbean lizards and with
barnacles
A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive ...
and their larvae along the California coast. In 2015, she wrote the fiction novel Ram-2050, a science-fiction retelling of the ''Ramayana.''
Research
Caribbean Anoles & Interspecific Competition
Roughgarden's early work in the 1970s and 80s helped to develop the ''
Anolis
''Anolis'' is a genus of anoles (), iguanian lizards in the family Dactyloidae, native to the Americas. With more than 425 species, it represents the world's most species-rich amniote tetrapod genus, although many of these have been proposed to ...
'' lizards of the Caribbean as an important model system for evolution and ecology. For example, she used she two-species enclosure experiments on two Caribbean islands to demonstrate increasing strength of interspecific competition as resource partitioning decreases: a central tenet for competition theory. The ''Anolis'' system thus provided an early example of an eco-evolutionary feedback, and with further development by
Jonathan Losos
Jonathan B. Losos (born December 7, 1961, in St. Louis County, Missouri) is an American evolutionary biologist and Herpetologist.
Life
Losos studied biology at Harvard University, from which he received a Bachelor's degree in 1984. Later on, in 19 ...
and others, has become an important example of
adaptive radiation
In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
.
Barnacles & Recruitment Limitation
After setting up a lab at the
Hopkins Marine Station Hopkins Marine Station is the marine laboratory of Stanford University. It is located ninety miles south of the university's main campus, in Pacific Grove, California (United States) on the Monterey Peninsula, adjacent to the Monterey Bay Aquarium ...
, Roughgarden sought to extend her approach of combining theoretical with field research by studying intertidal acorn
barnacle
A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in eros ...
s (''
Balanus
''Balanus'' is a genus of barnacles in the family Balanidae of the subphylum Crustacea.
This genus is known in the fossil record from the Jurassic to the Quaternary periods (age range: from 189.6 to 0.0 million years ago.). Fossil shells withi ...
'' and ''
Chthamalus
''Chthamalus'' (χθαμαλός, "flat" or "on the ground") is a genus of barnacles that is found along almost all non-boreal coasts of the northern hemisphere, as well as many regions in the southern hemisphere. These small barnacles have been ...
'' spp). Earlier work by
Joseph Connell
Joseph Hurd Connell FAA (5 October 1923 – 1 September 2020) was an American ecologist. He earned his MA degree in zoology at the University of California, Berkeley and his PhD at Glasgow University.This Week's Citation Classic. Garfield lib ...
Pisaster
''Pisaster'' (from Greek ', "pea", and ', "star"Cleveland P. Hickman et al., ''Integrated Principles of Zoology'' (St. Louis: Times Mirror / Mosby College Pub., 1984),p. 469) is a genus of Pacific sea stars that includes three species, ''P. b ...
'' seastars) and by competition, wherein dominant ''Balanus'' species displaced ''Chthamalus'' species to the high intertidal zones. Together with her student, Steve Gaines, Roughgarden showed that these interspecific interactions were most important in intertidal localities and communities with a high density of barnacles, such as those Connell and others had studied in
Scotland
Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to the ...
. At Hopkins in Central California, however, barnacle density was lower, and the amount of free space was best explained by periodic pulses of larval recruitment. With her student Sean Connolly, she then showed, through both empirical observation and modeling, that a latitudinal gradient in upwelling along the west coast of North America created very dense barnacle recruitment in the north (Oregon and Washington), where upwelling was weak, and very sparse barnacle recruitment in the south (California), where upwelling was strong. This in turn explained why field studies in the north had found interspecific interactions to be important, while her own field studies in the south had found larval recruitment to be most important for structuring intertidal populations. This deft synthesis helped to drive a paradigm shift in marine ecology which emphasized larval dispersal and recruitment dynamics over adult interactions and favored demographic models of populations open to larval recruits from distant localities, which dominated the field during the 1990s.
Criticism of sexual selection
Around the time of her transition, Roughgarden began to shift her research focus to Darwin’s theory of
sexual selection
Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
. In her 2004 book, ''Evolution’s Rainbow'', Roughgarden analyzes how biology can influence human sexuality and gender identity and explores the substantial diversity of mating systems and sexuality throughout the animal kingdom, with an eye toward understanding human sexual categories like gay, lesbian, and trans. In this book, and other articles around the same time, she offered criticism of sexual selection theory by providing examples of species that depart from its predictions (such as homosexual behavior in bonobos, elephants and lizards) as well as highlighting contradictions between population genetic theory and sexual selection theory. She also provided the beginnings of an alternative theory to sexual selection called social selection, which she describes as being focused on natural selection based on differential offspring production, where sexual selection is focused on differential mating success.
A 2006 article in the journal ''
Science
Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe.
Science may be as old as the human species, and some of the earliest archeological evidence for ...
'' with her student Erol Akçay, and Meeko Oishi, formally presented the theory of social selection in terms of game theory. Beyond the Nash Competitive Equilibrium (NCE) imported to evolutionary biology by
John Maynard Smith
John Maynard Smith (6 January 1920 – 19 April 2004) was a British theoretical and mathematical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics und ...
Nash Bargaining Solution Cooperative bargaining is a process in which two people decide how to share a surplus that they can jointly generate. In many cases, the surplus created by the two players can be shared in many ways, forcing the players to negotiate which division o ...
(NBS), which exists as an alternative to the NCE that is reached through negotiation. When playing in developmental time (as opposed to evolutionary time), a game player that stands to lose individual fitness at an NCE relative to its competitor may establish a threat point by promising to play a sub-optimal strategy. Through negotiation, for example via a side payment, the players can arrive at an NBS through playing mixed strategies across repeated games which thereby maximizes the fitness of the cooperative “team” (which consists of both players) rather than to any one player. Roughgarden et al. provide several examples of what such cooperative game play would look like in nature, and then define the evolutionary theory of social selection as one which considers such cooperative team games in the developmental tier as the primitive state, with sexual conflict as the derived state. They argue that social selection theory is mutually exclusive with the evolutionary theory of sexual selection, which treats sexual conflict as the primitive state and sexual cooperation as derived.
Following the ''Science'' paper, forty scientists produced ten critical letters stating that the article was misleading, that it contained misunderstandings and misrepresentations, that sexual selection accounted for all the data presented and subsumed Roughgarden's theoretical analysis, and that sexual selection explained data that her theory could not. Troy Day stated that "many people felt that this was completely shoddy science and poor scholarship, all motivated by a personal agenda". Roughgarden stated she was "not altogether surprised" by the volume of dissent and that her theory was not an extension of sexual selection theory.
Tim Clutton-Brock
Timothy Hugh Clutton-Brock (born 13 August 1946) is a British zoologist known for his comparative studies of the behavioural ecology of mammals, particularly red deer and meerkats. in which he concedes the point that males can engage in sexual selection on females, even in species where the operational sex ratio is biased towards males, stating: “Consequently, satisfactory explanations of the evolution of sex differences requires an understanding of the operation of sexual slection in females as well as males”. Nevertheless, Clutton-Brock concludes that sexual selection is a robust theoretical framework, without ever addressing the theoretical distinction in the polarity of intersexual cooperation and conflict highlighted by Roughgarden et al.
In her 2009 book ''The Genial Gene'', Roughgarden continues to build a case against sexual selection theory and to present
social selection
Social selection is a term used with varying meanings in biology.
Joan Roughgarden proposed a hypothesis called ''social selection'' as an alternative to sexual selection. Social selection is argued to be a mode of natural selection based on re ...
theory as an alternative. The book is titled as a response to the popular book, ''The Selfish Gene'', by
Richard Dawkins
Richard Dawkins (born 26 March 1941) is a British evolutionary biologist and author. He is an emeritus fellow of New College, Oxford and was Professor for Public Understanding of Science in the University of Oxford from 1995 to 2008. An ath ...
, which expounds what Roughgarden describes a “neo-Spencerian” view of nature “red in tooth and claw” in which competition and conflict dominate. In ''The Genial Gene'', after an initial section defining and attacking sexual selection, followed by a definition of social selection as one based on differences in offspring-production, rather than differences in mating success.
A second section is focused on the genetic basis of social selection. The first chapter addresses how sexual reproduction evolved in the first place, and makes the case for Roughgarden’s Portfolio hypothesis, which emphasizes that sexual reproduction creates genetic diversity through recombination, as opposed to the more commonly favored Muller’s Ratchet, which emphasizes that sex removes deleterious mutations through recombination. The second chapter explains the binary distribution of gamete types (sperm vs. egg) as a strategy to maximize gametic contact, rather than as a result of conflicting gametic strategies. A final chapter argues that hermaphroditism, rather than gonochorism, is the primitive state of sexuality.
The third section takes a two-tier approach to developing social selection theory. A behavioral tier focuses on game theory and Nash Bargaining Solutions as outlined in her 2006 paper. A second population genetic tier is then described that operates as a result of many replays of the behavioral tier. The book concludes by listing 26 phenomena that, according to Roughgarden, are explained differently by sexual selection theory and are better explained by social selection theory. She says that sexual selection theory derives from a view of natural behavior predicated on the selfish-gene concept, competition and deception, whereas the social-selection theory derives from teamwork, honesty, and genetic equality. As of 2012 she has continued to study if social selection as opposed to sexual selection is a more important driver of evolution for colonial species such as corals or perhaps humans.
Roughgarden's criticism of sexual selection has been rejected by the scientific community, and her papers on it have received few citations in scientific literature. In a 2019 interview, she stated that "Most biologists remain defensive of sexual selection theory”.
In 2013, Roughgarden funded a Catalysis meeting at the National Evolutionary Synthesis Center with the goal of debating and reviewing "the status of sexual selection studies and to indicate challenges and future directions". The group struggled to come to a consensus definition of sexual selection, but a subgroup offered a definition that for the first time explicitly differentiated fecundity selection for sexual selection ''sensu strictu''.
Theistic evolution
Roughgarden has written on the relationship between Christianity and science. Her book ''Evolution and Christian Faith: Reflections of an Evolutionary Biologist'' presents scripture passages that emphasize her belief that the Bible does not conflict with
evolutionary biology
Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life fo ...
and relates Christianity and evolution by asserting that all life is interconnected, as members of a faith community are connected. Roughgarden opposes
creationism
Creationism is the religious belief that nature, and aspects such as the universe, Earth, life, and humans, originated with supernatural acts of divine creation. Gunn 2004, p. 9, "The ''Concise Oxford Dictionary'' says that creationism is 't ...
and
intelligent design
Intelligent design (ID) is a pseudoscientific argument for the existence of God, presented by its proponents as "an evidence-based scientific theory about life's origins". Numbers 2006, p. 373; " Dcaptured headlines for its bold attempt to ...
Stonewall Book Award
The Stonewall Book Award is a set of three literary awards that annually recognize "exceptional merit relating to the gay/lesbian/bisexual/transgender experience" in English-language books published in the U.S. They are sponsored by the Rainbo ...
, 2005
*Center Fellow National Center for Ecological Synthesis and Analysis University of California (Santa Barbara), 1998
*Dinkelspiel Award for Undergraduate Teaching, Stanford University, in 1995
*Visiting Research Fellow at the Merton College, University of Oxford, in 1994
*Elected Fellow of the American Academy of Arts and Sciences in 1993
*Fellow of Guggenheim Foundation in 1985
*University Fellow, Stanford University in 1978
Roughgarden has served as associate editor of several
academic journals
An academic journal or scholarly journal is a periodical publication in which scholarship relating to a particular academic discipline is published. Academic journals serve as permanent and transparent forums for the presentation, scrutiny, and d ...
, including ''Philosophy and Theory in Biology'' (since 2008), ''
American Naturalist
''The American Naturalist'' is the monthly peer-reviewed scientific journal of the American Society of Naturalists, whose purpose is "to advance and to diffuse knowledge of organic evolution and other broad biological principles so as to enhance th ...
'' (1984–1989), ''
Oecologia
''Oecologia'' is an international peer-reviewed English-language journal published by Springer since 1968 (some articles were published in German or French until 1976). The journal publishes original research in a range of topics related to plant ...
'' (1979–1982), and ''
Theoretical Population Biology
''Theoretical Population Biology'' is a Peer review, peer-reviewed scientific journal covering research on Theoretical biology, theoretical aspects population biology in its widest sense, including mathematical modelling of populations, ecology, e ...
'' (1975–1986). She was the vice-chair and Chair of Theoretical Ecology Section of the
Ecological Society of America
The Ecological Society of America (ESA) is a professional organization of ecological scientists. Based in the United States and founded in 1915, ESA publications include peer-reviewed journals, newsletters, fact sheets, and teaching resources. I ...
during 2002–2003. She has served on the Nonprofit Organization Board for the Oceanic Society (San Francisco), the
EPA
The Environmental Protection Agency (EPA) is an independent executive agency of the United States federal government tasked with environmental protection matters. President Richard Nixon proposed the establishment of EPA on July 9, 1970; it be ...
Science Advisory Board Committee on Valuating the Protection of Ecological Systems and Services, and the science advisory boards of the Pacific Ocean Conservation Network (California), and the
Channel Islands National Marine Sanctuary
The Channel Islands National Marine Sanctuary is a sanctuary off the Pacific coast of Southern California. The National Marine Sanctuary program is under the administration of the National Oceanic and Atmospheric Administration.
Established ...
(Santa Barbara).
Selected bibliography
*
* Roughgarden, J. ''The Genial Gene: Deconstructing Darwinian Selfishness''. Hardcover ed. University of California Press, 2009
* Roughgarden, J. ''Evolution and Christian Faith: Reflections of an Evolutionary Biologist''. Hardcover ed. Washington, D.C.: Island Press, 2006.
* Roughgarden, J. ''Evolution's Rainbow: Diversity, Gender and Sexuality in Nature and People''. Berkeley CA: Univ. of California Press, 2004.
* Roughgarden, J. ''Primer of Ecological Theory''. 1st ed. Prentice Hall, 1997.
* Roughgarden, J. ''Anolis Lizards of the Caribbean: Ecology, Evolution and Plate Tectonics''. Hardcover ed. Oxford Univ. Press, 1995.
* Roughgarden, J, May, R. M., and Levin, S. A. (eds.). ''Perspectives in Ecological Theory''. Oxford Univ. Press, 1995.
*
* Roughgarden, J. ''Theory of Population Genetics and Evolutionary Ecology: an Introduction''. 1st ed. Prentice Hall, 1979.
*Simpson, Layne A. ''Gender and Society'', vol. 19, no. 3, 2005, pp. 425–426. ''JSTOR'', www.jstor.org/stable/30044605.
Bloggingheads.tv
Bloggingheads.tv (sometimes abbreviated "bhtv") is a political, world events, philosophy, and science video blog discussion site in which the participants take part in an active back and forth conversation via webcam which is then broadcast onl ...