Robert Paine at Mukkaw Bay, on the Olympic Peninsula in Washington. To understand the role of predatory starfish, he hurled them from an area and later returned to assess the sea life without them. Image credit: Kevin Schafer

It is with sadness that we share the news of the passing of Dr. Robert T. Paine III (M.S. Zoology 1958 and Ph.D.  Zoology 1961). Paine was an extraordinarily influential ecologist who coined the terms keystone species and trophic cascades, the strong, top-down effects that he and others discovered upon the removal or reintroduction of species.

His groundbreaking research, notably with starfish and sea otters, demonstrated for the first time what has become a hallmark of ecological study -- that keystone species and trophic cascades are integral to ecosystems and that rules of regulation govern the numbers and kinds of creatures in a community.

Paine’s years as a graduate student at the University of Michigan greatly influenced his research career. As a result of his interactions with the late ecologist Fred Smith, who taught a course on the natural history of freshwater invertebrates, Paine’s career took a U-turn from his original intention of geology and paleontology to the study of zoology, a field that truly captured his imagination. Smith became Paine’s advisor.

At the root of Paine’s academic family tree (illustrated on this page) lie his formative years in the U-M Department of  Zoology, a predecessor of the Department of Ecology and Evolutionary Biology. The tree depicts the vast number of ecologists taught by this stellar keystone among ecologists. We are honored to count Paine among our alumni.

Discover more about his career, legacy, and his time at the University of Michigan:

The Ecologist Who Threw Starfish, Nautilus, by Sean B. Carroll

RIP Bob Paine, A Keystone Among Ecologists, The Atlantic, by Ed Yong

Scientific families: Dynasty, Nature, by Ed Yong

Video: How Ecosystems Got a Keystone, Hakai magazine, directed by Jude Isabella and Adrienne Mason; produced by Gord More