Robert Treat "Bob" Paine III (April 13, 1933 – June 13, 2016) was 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 ...

who spent most of his career at the

University of Washington The University of Washington (UW, simply Washington, or informally U-Dub) is a public research university in Seattle, Washington. Founded in 1861, Washington is one of the oldest universities on the West Coast; it was established in Seattl ...

. Paine coined the keystone species concept to explain the relationship between '' Pisaster ochraceus'', a species of starfish, and '' Mytilus californianus'', a species of mussel.

Early life and education

Paine was born on April 13, 1933, and grew up in

Cambridge, Massachusetts Cambridge ( ) is a city in Middlesex County, Massachusetts, United States. As part of the Boston metropolitan area, the cities population of the 2020 U.S. census was 118,403, making it the fourth most populous city in the state, behind Boston ...

. He was fascinated by

biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary i ...

from a very young age.Natalie Hisdah
"An idea that spawned a legacy"
''Department of Biology eNews'',

, 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." After graduating 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 high ...

in 1954, he served in the

U.S. Army The United States Army (USA) is the land service branch of the United States Armed Forces. It is one of the eight U.S. uniformed services, and is designated as the Army of the United States in the U.S. Constitution.Article II, section 2, cl ...

, where he was the battalion gardener.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. Paine later entered graduate school at the

University of Michigan , mottoeng = "Arts, Knowledge, Truth" , former_names = Catholepistemiad, or University of Michigania (1817–1821) , budget = $10.3 billion (2021) , endowment = $17 billion (2021)As o ...

, intending to study

paleontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...

. Having taken some courses in

zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and ...

and

ecology 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 overl ...

at Michigan, his interests and studies changed after taking a course about freshwater

invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordat ...

taught by ecologist Frederick E. Smith. Upon graduating from the

, Paine completed a one-year postdoctoral fellowship at the Scripps Institution of Oceanography.Hillary Burgess (2013
"Diverse Introspectives: A conversation with Bob Paine"
''BioDiverse Perspectives; Graduate Students on Biodiversity Science'', 10 September 2013. Retrieved 15 June, 2016. In 1962, Paine joined the

, where he spent the rest of his career and became well known for his work.

Research career

Paine's doctoral research thesis was on the ecology of living brachipods.Robert T. Paine (1961
"The Life History and Population Dynamics of ''Glottidia Pyramidata'' (Brachiopoda)"
University of Michigan, PhD thesis. 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. In a noteworthyR.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." 1966 paper, Paine described a rocky

intertidal The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species ...

ecosystem in Makah Bay in

Washington state Washington (), officially the State of Washington, is a state in the Pacific Northwest region of the Western United States. Named for George Washington—the first U.S. president—the state was formed from the western part of the Washington ...

, where top predator species help maintain biodiversity. This led to his 1969 paper in which he proposed the keystone species concept. 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 Productivity is the efficiency of production of goods or services expressed by some measure. Measurements of productivity are often expressed as a ratio of an aggregate output to a single input or an aggregate input used in a production proces ...

. It has become a very popular concept in conservation biology. Paine also coined the term "

trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce t ...

" to describe the top-down effects that occur in ecosystems when an important species is introduced or removed.

Retirement and death

Paine retired in the late 1990s but continued to be active as a

professor emeritus ''Emeritus'' (; female: ''emerita'') is an adjective used to designate a retired chair, professor, pastor, bishop, pope, director, president, prime minister, rabbi, emperor, or other person who has been "permitted to retain as an honorary title ...

at the University of Washington.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." In 2000, he founded the Experimental and Field Ecology Fund to support graduate student research; the fund was renamed the Robert T. Paine Experimental & Field Ecology Endowed Fund to mark Paine's 80th birthday. As late as 2013, aged 79, Paine continued to make regular visits to

Tatoosh Island Tatoosh Island is a small island and small group of islands about offshore (northwest) of Cape Flattery, which is on the northwestern tip of the Olympic Peninsula in Washington. Tatoosh is the largest of a small group of islands also often refe ...

for research purposes. In 2013, he was awarded the

International Cosmos Prize The International Cosmos Prize was established in 1993, commemorating Expo '90 in Osaka, Japan. The objective of the prize was to develop the basic concept of Expo 90, "The Harmonious Coexistence between Nature and Mankind" and is awarded annuall ...

, including a cash prize equivalent to about US$408,000. Paine died from

acute myeloid leukemia Acute myeloid leukemia (AML) is a cancer of the myeloid line of blood cells, characterized by the rapid growth of abnormal cells that build up in the bone marrow and blood and interfere with normal blood cell production. Symptoms may inclu ...

, a type of

blood cancer Tumors of the hematopoietic and lymphoid tissues (American English) or tumours of the haematopoietic and lymphoid tissues (British English) are tumors that affect the blood, bone marrow, lymph, and lymphatic system. Because these tissues are all ...

, at the

Swedish Medical Center Swedish Health Services, formerly Swedish Medical Center, is the largest nonprofit health provider in the Seattle metropolitan area. It operates five hospital campuses (in the Seattle neighborhoods of First Hill, Cherry Hill and Ballard, and th ...

Seattle, Washington Seattle ( ) is a seaport city on the West Coast of the United States. It is the seat of King County, Washington. With a 2020 population of 737,015, it is the largest city in both the state of Washington and the Pacific Northwest region ...

, on June 13, 2016.Jennifer Ruesink (2016
"Prof. Emeritus Bob Paine (1933-2016)"
University of Washington, Department of Biology, 15 June 2016. Retrieved 16 June 2016. Phuong Le (2016

Phys.org Phys.org is an online science, research and technology news aggregator offering briefs from press releases and reports from news agencies (a form of journalism sometimes pejoratively called churnalism). The website also produces its own science jo ...

, 15 June 2016. Retrieved 16 June 2016.

Legacy

Paine's research—and the subsequent work of his students—has been very influential in the field of ecology, and he has been called a "giant" of the field. 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.

Ed Yong Edmund Soon-Weng Yong (born 17 December 1981) is a British-American science journalist. He is a staff member at '' The Atlantic'', which he joined in 2015. In 2021 he received a Pulitzer Prize for Explanatory Reporting for a series on the COVI ...

wrote that "by encouraging independence and prizing fieldwork, Paine mentored an entire generation of superstar ecologists." Paine's former students and postdocs include Paul Dayton, Bruce Menge and

Jane Lubchenco Jane Lubchenco (born December 4, 1947) is an American environmental scientist and marine ecologist who teaches and conducts research at Oregon State University. Her research interests include interactions between the environment and human well-be ...

. Paine and his work are featured prominently in the 2018 documentary film '' The Serengeti Rules''.

Recognition

*Vice-President,

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 ...

, 1977–1978 *President, Ecological Society of America, 1979–1980 *

Robert H. MacArthur Award The Robert H. MacArthur Award is a biennial prize given by the Ecological Society of America to ecologists for their pivotal contributions to their field. The acceptance speeches of many recipients have been given at the annual meeting of the soci ...

, Ecological Society of America, 1983Robert H. MacArthur Award

, 30 January 2014. Retrieved 15 June 2016. *Elected to

The National Academy of Sciences The National Academy of Sciences (NAS) is a United States nonprofit, non-governmental organization. NAS is part of the National Academies of Sciences, Engineering, and Medicine, along with the National Academy of Engineering (NAE) and the Nati ...

, 1986 *

Sewall Wright Award The Sewall Wright Award is given annually by the American Society of Naturalists to a "senior-level" and active investigator making fundamental contributions the conceptual unification of the biological sciences. The award was established in 1991 a ...

, 1996 *

American Society of Naturalists The American Society of Naturalists was founded in 1883 and is one of the oldest professional societies dedicated to the biological sciences in North America. The purpose of the Society is "to advance and diffuse knowledge of organic evolution and o ...

Honorary Lifetime Membership Award, 2009 *

, 2013Katherine Long (2013
"Retired UW prof wins $408,000 for groundbreaking ecology work"
''Seattle Times'', 2 August 2013. Retrieved 15 June 2016.

References

Notes

Citations

{{DEFAULTSORT:Paine, Robert T. 1933 births 2016 deaths University of Washington faculty 20th-century American zoologists Members of the United States National Academy of Sciences University of Michigan alumni Harvard University alumni People from Cambridge, Massachusetts Fellows of the Ecological Society of America American ecologists Deaths from cancer in Washington (state) Deaths from acute myeloid leukemia