April 13, 1933 – June 13, 2016
Dr. Robert (Bob) Paine holds iconic status within ecology, and his death brings both immense loss and a reminder that he will always be with us through his shaping of ecological ideas and practices.
He is justly famous for coining the term “keystone species” in reference to predators whose effects indirectly maintain the diversity, structure, and functioning of ecological communities. Like the keystone in an arch, these species hold a system together. Bob’s field study of a keystone species is so familiar as a textbook example that it hardly needs repeating: When he removed ochre sea stars from a rocky shoreline, mussels began to monopolize space, halving the number of species attached to the rocks. The description of the rocky shore food web, originally published in 1966 (with the keystone term appearing in a subsequent paper), has been cited several thousand times and reprinted as one of the key papers in the development of the discipline in “Foundations of Ecology” (1991). The global value of understanding how predators shape ecosystems comes to life in a recent video, including recollections of and interviews with Bob.
Bob helped rescue the keystone species concept several decades after its inception, when its application to any species considered for conservation had diluted its original meaning. The effort generalized the keystone term to encompass strong single-species impacts that were out of proportion to the species’ abundance. “The Importance of Species” book (2002) arose from a retirement tribute to Bob, highlighting his commitment to learning about strong species interactions. In recent collaborations, he has pointed out the shambles of structure and function resulting from trophic downgrading, or the loss of top predators from ecosystems.
Bob also transformed how ecology was carried out, ushering in to the last half of the 20th century an era of field experimentation rather than relying primarily on observations of the environment. The drawback of observations is that patterns or changes can happen for any number of reasons, whereas an experiment tests specific, plausible drivers. Bob’s commitment to field experimental ecology permeates his book, “Marine Rocky Shores and Community Ecology: An Experimentalist’s Perspective” (1994), where he wrote of “experimental manipulation as the surest means of both testing and rejecting hypotheses and generating novel insights.”
Bob was recognized internationally for his contributions to ecology, including eminent ecologist to deliver the Tansley Lecture to the British Ecological Society (1979), the MacArthur Award for lifetime achievement from the Ecological Society of America (1983), the ECI prize from the Ecology Institute in Germany (1989), and the International Cosmos Prize (2013). He was elected to the National Academy of Sciences in 1986. The accompanying explanations for these awards often make no mention of keystone species, but emphasize other foundational contributions to ecological ideas on food webs, patch dynamics, non-equilibrial nature of communities, or implications for biodiversity conservation.
How can one person contribute so many game-changing ideas? By his own assessment, these did not emerge from the intellectual milieu at the time, or from theoretical explorations, but from careful examination of living systems. Scientifically, he was inspired by nature.
From the start, Bob’s wooded neighborhood near Cambridge, Massachusetts was a paradise for a free-roaming, curious child. He was a self-described “boy birder” with a keen eye for detail and penchant for the outdoors. As he grew up, this study and care of nature became central elements of his vocation. He received his bachelors degree from Harvard University (1954) and PhD from the University of Michigan (1961). Bob joined the faculty of the Department of Zoology at the University of Washington in 1962, where he remained throughout his career, including eight years as chair. Even after reaching emeritus status, he maintained an active research program at his long-term study site on Tatoosh Island, among the best- studied ecological systems on the planet as recorded in his chapter of “The Ecology of Place” (2011, with C. Pfister and J.T. Wootton). Work on Tatoosh has been carried out with the permission of the Makah Tribe, for whom the island is part of their cultural home.
The outer coast of Washington, of which Tatoosh is the western-most gem, reveals stunning pattern and rich biodiversity at low tide. On simple two-dimensional surfaces, where different species appear with small changes in inundation time, the wave-exposed rocky intertidal zone became Bob’s natural laboratory. 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 home-made steps. Every two weeks during the summer, 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. Bob wrote his observations in bold strokes on plastic slates, each one then run through a copier for transfer to paper and permanent archiving by being taped into a leather-bound book. This series is now up to more than 20 volumes, containing decades of records of weather, bird lists, contents of the wrack line on the beach, as well as experimental results. As each tide rose and chased ecologists from their intertidal studies, Bob doled out a measure of box wine into a mug and sat down with his current scientific family to talk about what each had seen and learned.
Generations of students and colleagues have been touched directly and indirectly by Bob’s ideas. He instilled a love of natural history in those who shared time with him in the field, including field courses which he led not just at the University of Washington but worldwide. He held deep respect for nature and an esthetic appreciation of patterns, while nurturing the curiosity and interventions to find out how complex systems work. He remained true to his ideas that consumers – predators, grazers, herbivores, everything that eats – shape the world. In his honor, we can always ask, with a gleam in the eye and intent to spark a spirited discussion, “What would have happened in that study if the consumers were caged out?” He leaves a lasting legacy as a keystone ecologist, whose contributions – disproportionate even to his imposing biomass – influence the diversity of ideas about how to study and interpret the complex systems of the living earth.
This spring, I had the pleasure of leading a graduate seminar on modern ecological game-changers. Bob regularly weighed in by email, though he couldn’t attend in person. The “game-changer” topic had been chosen prior to Bob’s cancer diagnosis, but ended up providing a weekly opportunity for me to reflect on him. I was reminded of the graduate seminars that he organized himself, decades ago, which the students affectionately dubbed “Paine-bull” to reflect the exchange of perspectives, sometimes deliberately extreme to draw out debate. I was also reminded of Bob’s career-long commitment to graduate mentoring, which has grown a rich extended family of successful ecologists in their own right. Bob always held to the opinion that individual efforts were more cost-effective than Big Science, and that graduate students should practice independent research, rather than be cogs in bigger wheels. A perfect example of this opinion was his establishment of an endowed fund in Biology to support graduate research, re-named to include his name on the occasion of his 80th birthday: the Robert T. Paine Experimental and Field Ecology Award. This spring, the “game-changer” seminar raised an unexpected challenge of trying to figure out, given the three-year time window to which we had limited ourselves, what was really going to transform ecology, rather than be a weird splash or a dead end. So I asked Bob: how did you know that you had contributed a game-changer? He told me that a couple of months after his 1966 paper was published, he had received a letter from Robert MacArthur, the most prominent American ecologist at the time and a strict adherent to community structure driven by competition. MacArthur had read about sea stars as predators maintaining the diversity of rocky shores and was willing to admit, in writing to Bob: “This changes everything.”
Indeed it has. Thanks, Bob, for everything.
Bob's strong and loving daughters have concurred with Bob's own wishes to support graduate students, and that there would be no better place for memorial gifts than to the Paine Experimental and Field Ecology Endowed Fund in Biology at the University of Washington.
Sincerely, Jennifer Ruesink
Gifts in honor of Bob can be directed to the Robert T. Paine Experimental & Field Ecology Endowed Fund. Please make checks out to the University of Washington, with "Paine endowment" on the memo line. For questions, email <firstname.lastname@example.org> or call 206.685.2185
From Bob Paine’s Family:
We will be forever grateful for the outpouring of sympathy and support we have received from the University of Washington, and dad’s friends and many other professional and personal associates during the ordeal of his illness and passing. And our thanks go out to Swedish Hospital, Group Health, and the doctors and caregivers who provided the wonderful care that helped Dad through this difficult process and eased his pain.
This has been difficult for all concerned, and especially for his family. All of us are grateful that so many of Dad’s closest companions from his long career as a dedicated biologist traveled from places near and far to his bedside to show their loving support. All these visits, letters, and expressions of affection were enjoyed and appreciated by him – and all of us – gave us peace and closure through this ordeal.
Many thanks to you all!