RIP Bob Paine, A Keystone Among Ecologists

Bob Paine's academic family tree (Marian Kohn)
Editor’s Note: This article previously appeared in a different format as part of The Atlantic’s Notes section, retired in 2021.

I’m deeply saddened to learn that Bob Paine, a giant of ecology, passed away yesterday. You may not know his name, but you almost certainly know the ideas that he pioneered.

Back in 1963, Paine began prying ochre starfish off a rocky beach in Washington and hurling them into the sea. After a year, the mussels that the starfish would normally have eaten had overrun the beach, turning a wonderland of limpets, anemones, and barnacles into a monoculture of black gaping shells.

The experiment was ground-breaking. It showed that not all species are equal, and that some—like the starfish—are secret lynchpins of the natural world. Their absence can ripple outwards, triggering the rise and fall of connected species and can even reshape the landscape. For example, when sea otters vanish, the sea urchins they eat transform lush forests of kelp into desolate barrens, dooming the fish, crabs, and other animals that once lived there. Paine called these ripples “trophic cascades”, and he billed the animals behind them—the starfish, otters, and others—as “keystone species”, after the central stone that stops an arch from collapsing. These concepts are so familiar today that we take them for granted, but we didn’t always know about them. We only do because of Paine.

I met Paine in 2012 to write a story for Nature about his legacy. Like the starfish he studied, he has also been disproportionately influential. By encouraging independence and prizing fieldwork, Paine mentored an entire generation of superstar ecologists. They, their students, and their student’s students have changed our understanding of the natural world. You can read about the entire dynasty here.

Despite his influence, Paine struck me as unusually humble. He was certainly opinionated and occasionally cantankerous, but also warm, good-humoured and deeply self-effacing. He loathed the limelight as much as he loved nature. Consider this snippet from a departmental email from Toby Bradshaw from the University of Washington, in which he announced Paine’s passing:

“While I doubt that anyone has ever used the words “shy” or “reticent” to describe Bob, he kept his incredibly generous philanthropy towards our department very quiet, though he told me that he was deeply proud of all the graduate students who received the Experimental and Field Ecology Award from its anonymous donor.  It took the persuasive talents of Jennifer Ruesink and Ray Huey to convince Bob that the students would be best served by having his endowment renamed the Robert T. Paine Experimental and Field Ecology Award, on the occasion of Bob’s 80th birthday.”

That was Paine. He didn’t care for glory, reflected or otherwise. He just wanted smart people to win. And thanks to him, many did.

RIP, Bob. It was a pleasure to have met you.