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NY Times, June 18 2016
Robert Paine, Ecologist Who Found ‘Keystone Species,’ Dies at 83
By SAM ROBERTS
Robert Paine, a groundbreaking, hands-on ecologist who found that
removing what he called a “keystone species” from an environment could
profoundly affect the fortunes of neighboring species, died on Monday in
Seattle. He was 83.
The cause was acute myeloid leukemia, his daughter Anne Paine said.
Dr. Paine demonstrated in his field work that certain species exert a
disproportionate impact on their ecosystems and that their elimination —
as a result of climate change, pollution or some other natural or
man-made factors — can produce unexpected and far-reaching consequences
for the local environment.
A teacher and researcher at the University of Washington for 36 years,
Dr. Paine propounded his keystone theory in 1966 after studying ochre
starfish, or sea stars, as they preyed on the mussel population along
the rocky shore of Makah Bay, on the tip of the Olympic Peninsula in
Washington State.
After he pried the starfish from rocks with a crowbar and hurled them
into the sea, the mussels proliferated along the shore, displacing algae
and limpets.
He found a similar chain reaction — or “trophic cascade,” as he called
it — when sea otters vanished or were removed from an environment
because of fur trading, pollution or marine predators. With the otters
gone, sea urchins, which the otters had preyed upon, were now free to
gobble up a larger share of kelp — food that would otherwise have
sustained fish and crabs.
He identified the predator starfish and the otters as keystone species,
taking the name from the wedge-shaped apex of an arch that keeps it from
collapsing.
Dr. Paine, who had a passion for field work, conducted much of his own
research on Tatoosh Island, an uninhabited rocky outcropping less than a
mile off Cape Flattery, on the Olympic Peninsula. He discovered the spot
in 1967 on a salmon-fishing trip.
Simon Levin, an ecologist and professor at Princeton University, wrote
in an email that Dr. Paine’s “influence cannot be overestimated,”
particularly his “notion that to understand systems one had to perturb
them.”
“He helped make ecology an experimental science,” Professor Levin wrote.
Dr. Paine quickly achieved a stature in the scientific community that
matched his 6-foot-4 frame. Several months after he published his
seminal paper on keystone species in the journal American Naturalist in
1966, he received a letter from Robert H. MacArthur, a leading ecologist
at Princeton.
“This changes everything,” Professor MacArthur wrote.
Robert Treat Paine III was born in Cambridge, Mass., on April 13, 1933.
His father, also named Robert, was a curator at the Museum of Fine Arts
in Boston and a descendant of Robert Treat Paine, a Massachusetts lawyer
who was a signer of the Declaration of Independence. His mother, the
former Barbara Birkhoff, was a writer and photographer.
His marriage to the former Alice E. Coleman ended in divorce. In
addition to his daughter Anne, he is survived by two other daughters,
Susan Paine and Nancy Paine; five grandchildren; a great-granddaughter;
and a brother, Garrett.
“All my early childhood memories involve biology,” Dr. Paine said in an
online interview with the University of Washington’s biology department.
“I remember sitting in the dirt driveway when I was around 2½ years old
and watching ants. I was utterly fascinated with nature from a very
young age.”
That fascination, along with an interest in birding, propelled him
toward a study of fossils as a paleontologist and geologist. After
graduating with a bachelor’s degree from Harvard University in 1954, he
served in the Army, then pursued graduate studies at the University of
Michigan.
But at Michigan he shifted his focus, from extinct to animate objects,
after taking a revelatory course on freshwater invertebrates given by
the naturalist Frederick E. Smith, a faculty member. Inspired, he went
on to earn a master’s degree and a doctorate in zoology.
He joined the faculty of the University of Washington in 1962 and was
promoted to professor in 1971. He retired in 1998, but returned to
campus regularly and established an endowment to support graduate research.
Dr. Paine preferred field work, as Jennifer Ruesink, a biology professor
at the University of Washington, wrote in a tribute to him on the
university’s website:
“Working on a remote island was not easy: leaps of faith across surge
channels, slippery algae that could take down the most sure-footed,
boats overturned and scientists bodily moved by rogue waves, all
supplies hefted from the beach to the top of the island via a hundred
homemade steps.
“Every two weeks during the summer,” she added, “when the tides reached
their extreme lows, and at longer intervals in winter for tides in the
dark, Tatoosh Island had its vital signs checked and challenged.”
Dr. Paine found great value in deliberately disrupting an environment in
the interest of science, as he did on Makah Bay, when he threw the
predator starfish into the waves, away from the shore. “Experimental
manipulation is not only more interesting, it’s much more fun,” he told
The Seattle Times in 2013. “And getting results you can interpret, if
you test an idea — that’s what science is all about.”
He continued to teach well into his later years and produced a dynasty
of ecologists. “All my students were smarter than me,” he was quoted as
saying in a 2013 profile in The Atlantic magazine, “but just less
knowledgeable.”
One former student, Peter Kareiva, the director of the Institute of the
Environment and Sustainability at the University of California, Los
Angeles, wrote of Dr. Paine in an email: “He insisted on experiments in
ecology at a time when others were content with simply explaining
patterns. I think he turned ecology from quantitative natural history
into a modern science.”
Later in his career, with his concerns about the impact of human
activity on the environment growing, Dr. Paine came to recognize the
most crucial keystone species of them all.
“Humans are certainly the overdominant keystones,” he said earlier this
year in the science journal Nautilus, “and will be the ultimate losers
if the rules are not understood and global ecosystems continue to
deteriorate.”
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