Robert T. Paine (zoologist)

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Robert T. Paine
Born(1933-04-13)April 13, 1933
DiedJune 13, 2016(2016-06-13) (aged 83)
Alma materHarvard University
University of Michigan
Known forkeystone species concept[1]
AwardsSewall Wright Award (1996)
National Academy of Sciences
International Cosmos Prize (2013)
Scientific career
InstitutionsUniversity of Michigan
University of Washington
Harvard University
Scripps Institution of Oceanography
ThesisThe Life History and Population Dynamics of Glottidia Pyramidata (Brachiopoda) (1961)
Doctoral studentsPaul Dayton[1]
Bruce Menge[1]
Jane Lubchenco[1]
Anne Salomon

Robert Treat "Bob" Paine III (April 13, 1933 – June 13, 2016) was an American ecologist who spent most of his career at the University of Washington. Paine coined the keystone species[1][2][3] concept to explain the relationship between Pisaster ochraceus, a species of starfish, and Mytilus californianus, a species of mussel.[4]

Early life and education[edit]

Paine was born on April 13, 1933,[5] and grew up in Cambridge, Massachusetts.[6] He was fascinated by biology from a very young age.[7] After graduating from Harvard University in 1954, he served in the U.S. Army, where he was the battalion gardener.[8] Paine later entered graduate school at the University of Michigan, intending to study paleontology.[6] Having taken some courses in zoology and ecology at Michigan, his interests and studies changed after taking a course about freshwater invertebrates taught by ecologist Frederick E. Smith.[6][8] Upon graduating from the University of Michigan, Paine completed a one-year postdoctoral fellowship at the Scripps Institution of Oceanography.[9] In 1962, Paine joined the University of Washington, where he spent the rest of his career and became well known for his work.[8]

Research career[edit]

Paine's doctoral research thesis was on the ecology of living brachipods.[10] As a postdoctoral fellow, he worked on the history and energetics of opisthobranchs (marine gastropods). Much of Paine's work at the University of Washington focused on the organization of marine communities. It was here that much of his research on keystone species occurred.[8][11]

In a noteworthy[12] 1966 paper, Paine described a rocky intertidal ecosystem in Makah Bay in Washington state, where top predator species help maintain biodiversity.[13] This led to his 1969 paper in which he proposed the keystone species concept.[14][15][16] This concept states that an ecosystem may experience a dramatic shift if a keystone species is removed, even though that species was a small part of the ecosystem by measures of biomass or productivity. It has become a very popular concept in conservation biology.[17] Paine also coined the term "trophic cascade" to describe the top-down effects that occur in ecosystems when an important species is introduced or removed.[6][11]

Retirement and death[edit]

Paine retired in the late 1990s but continued to be active as a professor emeritus at the University of Washington.[18] In 2000, he founded the Experimental and Field Ecology Fund to support graduate student research;[7] the fund was renamed the Robert T. Paine Experimental & Field Ecology Endowed Fund to mark Paine's 80th birthday.[1][5] As late as 2013, aged 79, Paine continued to make regular visits to Tatoosh Island for research purposes.[1][A]

In 2013, he was awarded the International Cosmos Prize, including a cash prize equivalent to about US$408,000.[19] Paine died from acute myeloid leukemia, a type of blood cancer, at the Swedish Medical Center in Seattle, Washington, on June 13, 2016.[5][11] [20]


Paine's research—and the subsequent work of his students—has been very influential in the field of ecology,[1] and he has been called a "giant" of the field.[20][21] Paine's research helped popularise field manipulation experiments, sometimes called "kick-it-and-see ecology", at a time when field ecologists tended only to observe natural ecosystems.[1] Ed Yong wrote that "by encouraging independence and prizing fieldwork, Paine mentored an entire generation of superstar ecologists."[21] Paine's former students and postdocs include Paul Dayton, Bruce Menge and Jane Lubchenco.[1] Paine and his work are featured prominently in the 2018 documentary film The Serengeti Rules.[22]




  1. ^ Close to Cape Flattery, "the northwesternmost point of the continental United States", the island is owned by the Makah Indian tribe. They granted him permission to conduct his research, with the only restriction being that he not ‘‘mess with the graves.’’[11]


  1. ^ a b c d e f g h i j Yong, E. (2013). "Scientific families: Dynasty. Bob Paine fathered an idea — and an academic family — that changed ecology". Nature. 493 (7432): 286–289. doi:10.1038/493286a. PMID 23325190.
  2. ^ "Keystone Species Hypothesis". University of Washington. Archived from the original on 2011-01-10. Retrieved 2011-02-03.
  3. ^ "Scientists Adopt Tiny Island as a Warming Bellwether". New York Times. 6 Oct 2012. Retrieved 2012-10-25.
  4. ^ Stolzenberg, William (2008). Where the Wild Things Were: Life, death and ecological wreckage in a land of vanishing predators. Bloomsbury USA. ISBN 978-1-59691-299-1.
  5. ^ a b c Jennifer Ruesink (2016) "Prof. Emeritus Bob Paine (1933-2016)" University of Washington, Department of Biology, 15 June 2016. Retrieved 16 June 2016.
  6. ^ a b c d Carroll, Sean B. (10 March 2016). "The Ecologist Who Threw Starfish". Nautilus. Retrieved 11 March 2016.
  7. ^ a b Natalie Hisdahl "An idea that spawned a legacy" Archived 2015-11-14 at the Wayback Machine Department of Biology eNews, University of Washington, Summer 2013. Retrieved 15 June, 2016. "All my early childhood memories involve biology. I remember sitting in the dirt driveway when I was around two-and-a-half years old and watching ants—I was utterly fascinated with nature from a very young age."
  8. ^ a b c d e f R.B. Root (1979) "Robert T. Paine, President: 1979–1980" Bulletin of the Ecological Society of America, 60(3): 156–157 (September 1979). Retrieved 15 June 2016.
  9. ^ Hillary Burgess (2013) "Diverse Introspectives: A conversation with Bob Paine" BioDiverse Perspectives; Graduate Students on Biodiversity Science, 10 September 2013. Retrieved 15 June, 2016.
  10. ^ Robert T. Paine (1961) "The Life History and Population Dynamics of Glottidia Pyramidata (Brachiopoda)" University of Michigan, PhD thesis.
  11. ^ a b c d Langer, Emily (June 17, 2016). "Bob Paine, 83, ecologist whose work identified keystone species". Washington Post. Retrieved June 17, 2016.
  12. ^ R.B. Root (1979) "Robert T. Paine, President: 1979–1980" Bulletin of the Ecological Society of America, 60(3): 156–157 (September 1979). Retrieved 15 June 2016. "This paper, which has been widely cited and reprinted, was one of the first clear proofs of a phenomenon which [...] appears to be an important organizing factor in several ecosystems."
  13. ^ Paine, R. T. (1966). "Food Web Complexity and Species Diversity". The American Naturalist. 100 (910): 65–75. doi:10.1086/282400. S2CID 85265656.
  14. ^ Paine, R. T. (1969). "A Note on Trophic Complexity and Community Stability". The American Naturalist. 103 (929): 91–93. doi:10.1086/282586. S2CID 83780992.
  15. ^ a b "Honorary Lifetime Membership Award: Robert T. Paine". The American Naturalist. 174 (3): iii–. 2009. doi:10.1086/605920. PMID 19653846. S2CID 205992377.
  16. ^ a b c Wilbur, H. M. (1998). "1996 Sewall Wright Award: Robert T. Paine". The American Naturalist. 151 (1): i–t. doi:10.1086/513833. PMID 18811418. S2CID 39717819.
  17. ^ Mills, L. Scott; Soulé, Michael E.; Doak, Daniel F. (1993). "The Keystone-Species Concept in Ecology and Conservation". BioScience. 43 (4): 219–224. doi:10.2307/1312122. JSTOR 1312122.
  18. ^ Katherine Long (2013) "Retired UW prof wins $408,000 for groundbreaking ecology work" Seattle Times, 2 August 2013. Retrieved 15 June 2016. "Paine, 80, has been retired for 15 years, yet he still shows up on campus daily to work out of his cluttered basement office, in Kincaid Hall, writing and contributing to research papers."
  19. ^ a b Katherine Long (2013) "Retired UW prof wins $408,000 for groundbreaking ecology work" Seattle Times, 2 August 2013. Retrieved 15 June 2016.
  20. ^ a b Phuong Le (2016) "Bob Paine, ecologist who introduced 'keystone species,' dies", 15 June 2016. Retrieved 16 June 2016.
  21. ^ a b Yong, Ed (14 June 2016). "RIP Bob Paine, A Keystone Among Ecologists". The Atlantic. Retrieved 2016-06-14.
  22. ^ John DeFore (2019) "'The Serengeti Rules': Film Review" The Hollywood Reporter. Published May 8, 2019. Accessed June 6, 2019.
  23. ^ Robert T. Paine (1983) "Ecological Determinism in the Competition for Space: The Robert H. MacArthur Award Lecture" Ecology, 65: 1339-1348.
  24. ^ Robert H. MacArthur Award, Ecological Society of America, 30 January 2014. Retrieved 15 June 2016.