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

ecologist Ecology () is the natural science of the relationships among living organisms and their environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely re ...

who spent most of his career at the

University of Washington The University of Washington (UW and informally U-Dub or U Dub) is a public research university in Seattle, Washington, United States. Founded in 1861, the University of Washington is one of the oldest universities on the West Coast of the Uni ...

. Paine coined the

keystone species A keystone species is a species that has a disproportionately large effect on its natural environment relative to its abundance. The concept was introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in main ...

concept to explain the relationship between ''

Pisaster ochraceus ''Pisaster ochraceus'', generally known as the purple sea star, ochre sea star, or ochre starfish, is a common seastar found among the waters of the Pacific Ocean. Identified as a keystone species, ''P. ochraceus'' is considered an important ind ...

'', a species of

starfish Starfish or sea stars are Star polygon, star-shaped echinoderms belonging to the class (biology), class Asteroidea (). Common usage frequently finds these names being also applied to brittle star, ophiuroids, which are correctly referred to ...

, and ''

Mytilus californianus The California mussel (''Mytilus californianus'') is a large edible mussel, a marine bivalve mollusk in the family Mytilidae. This species is native to the west coast of North America, occurring from northern Mexico to the Aleutian Islands of ...

'', a species of

mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and Freshwater bivalve, freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other ...

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. It is a suburb in the Greater Boston metropolitan area, located directly across the Charles River from Boston. The city's population as of the 2020 United States census, ...

. He was fascinated by

biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...

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 university, private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the History of the Puritans in North America, Puritan clergyma ...

in 1954, he served in the

U.S. Army The United States Army (USA) is the primary land service branch of the United States Department of Defense. It is designated as the Army of the United States in the United States Constitution.Article II, section 2, clause 1 of the United Stat ...

, where he was the battalion gardener.R.B. Root (1979
"Robert T. Paine, President: 1979–1980"
''

Bulletin of the Ecological Society of America The Ecological Society of America (ESA) is a professional organization of ecology, ecological scientists. Based in the United States and founded in 1915, ESA publications include peer-reviewed journals, newsletters, fact sheets, and teaching reso ...

'', 60(3): 156–157 (September 1979). Retrieved 15 June 2016. Paine later entered graduate school at the

University of Michigan The University of Michigan (U-M, U of M, or Michigan) is a public university, public research university in Ann Arbor, Michigan, United States. Founded in 1817, it is the oldest institution of higher education in the state. The University of Mi ...

, intending to study

paleontology Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure ge ...

. Having taken some courses in

zoology Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...

and

ecology Ecology () is the natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...

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

invertebrates Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...

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

, Paine completed a one-year postdoctoral fellowship at the

Scripps Institution of Oceanography Scripps Institution of Oceanography (SIO) is the center for oceanography and Earth science at the University of California, San Diego. Its main campus is located in La Jolla, with additional facilities in Point Loma. Founded in 1903 and incorpo ...

.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

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 or foreshore is the area above water level at low tide and underwater at high tide; in other words, it is the part of the littoral zone within the tidal range. This area can include several types of habitats with various sp ...

ecosystem in

Makah Bay Makah Bay is a bay in Clallam County, Washington, United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. stat ...

Washington state Washington, officially the State of Washington, is a state in the Pacific Northwest region of the United States. It is often referred to as Washington State to distinguish it from the national capital, both named after George 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 Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...

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 proce ...

. It has become a very popular concept in

conservation biology Conservation biology is the study of the conservation of nature and of Earth's biodiversity with the aim of protecting species, their habitats, and ecosystems from excessive rates of extinction and the erosion of biotic interactions. It is an i ...

. 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 redu ...

" 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/Emerita'' () is an honorary title granted to someone who retirement, retires from a position of distinction, most commonly an academic faculty position, but is allowed to continue using the previous title, as in "professor emeritus". ...

at the University of Washington.Katherine Long (2013
"Retired UW prof wins $408,000 for groundbreaking ecology work"
''

Seattle Times ''The Seattle Times'' is an American daily newspaper based in Seattle, Washington. Founded in 1891, ''The Seattle Times'' has the largest circulation of any newspaper in the state of Washington and the Pacific Northwest region. The Seattle Time ...

'', 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 ...

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 Humankind" and is awarded by t ...

, 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 haematopoiesis, normal blood cell production. Sympt ...

, 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 al ...

, at the

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

Seattle, Washington Seattle ( ) is the List of municipalities in Washington, most populous city in the U.S. state of Washington (state), Washington and in the Pacific Northwest region of North America. With a population of 780,995 in 2024, it is the List of Unit ...

, 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 which re-publishes press releases and stories from news agencies (a business model known as churnalism). In 2012, PhysOrg.com was changed to Phys.org. , Phys.org was posting ...

, 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 Americans, British-American science journalist and author. In 2021, he received a Pulitzer Prize for Explanatory Reporting for a series on the COVID-19 pandemic. He is the author of two ...

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- ...

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

The Serengeti Rules ''The Serengeti Rules'' is a 2018 American documentary film directed by Nicolas Brown, and based on the book by Sean B. Carroll. The film explores the discoveries of five pioneering scientists— Tony Sinclair, Mary E. Power, Bob Paine, John Te ...

''.

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 soc ...

, 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 Natio ...

, 1986 *

Sewall Wright Award ASN Award for Distinguished Achievement in the Conceptual Unification of the Biological Sciences, previously known as the Sewall Wright Award, is given annually by the American Society of Naturalists to a "senior-level" and active investigator mak ...

, 1996 *

American Society of Naturalists The American Society of Naturalists (ASN) 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 evoluti ...

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 Scientists from Cambridge, Massachusetts Fellows of the Ecological Society of America American ecologists Deaths from leukemia in Washington (state) Deaths from acute myeloid leukemia