“All Animals Weren’t Created Equal” Robert T. Paine

1. “All Animals Weren’t Created Equal”Robert T. Paine Jenessa Kay April 23, 2013 Community Ecology Nature Education, 2012

2. Biographical Info • Born in Cambridge, MA – April 13,1933 • Happy belated birthday! • AB - Harvard University, 1954 • Army Battalion Gardener • PhD - University of Michigan, 1961 • Post-doc, Scripps Institute of Oceanography • Professor of Zoology, University of Washington 1962 – 1998 • Current Professor emeritus of Zoology, UW Kinne (ed.), 1994

3. Awards and Recognition • 1979-80 – President of Ecological Society of America • 1983 – MacArthur Award (ESA) • 1989 – Excellence in Ecology Prize (Ecology Inst. in Oldendorf am Luhe, Germany • 1997 – Sewall Wright Award (Society of American Naturalists) • 2000 – Eminent Ecologist Award Scientific American, 2010

4. Cited by 776 Cited by 3652 Cited by 1178 Cited by 920 BioScience, Vol. 46, No. 8 (Sep., 1996) Cited by 1017

5. Research Overview • Predation Hypothesis and Competitive Hierarchies • Food Web Interactions and Trophic Cascades • Quantifying interaction strengths and patch dynamics • Keystone Species Concept marinebio.net Why don’t we have monocultures of good competitors??

6. The New York Times, 2012 Kevin Schafer

7. Study Location • Makah Bay – mainland WA • Tatoosh Island - 0.5 miles offshore from Cape Flattery • Part of Makah Reservation • Longest ongoing study of a single area by the same scientist in the U.S. makah.com Superstock.com

8. Tatoosh Island legendsofamerica.com wikimedia.com history.noaa.gov (1943)

9. “Whether I was stupid or foolhardy, I spent my first ten years in the intertidal in sneakers – cheap as possible.” scientificamerican.com “I have too good peripheral circulation.”

10. The New York Times, 2012

11. “His intellectual presence is so commanding that his physical presence hardly registers.” naturalhistoriesproject.org

12. “If you ask him a question…you feel the weight of an encyclopedic knowledge of scientific and natural history gathering behind his response.” naturalhistoriesproject.org

13. “What would happen if we removed the top predator from an ecosystem?” naturalhistoriesproject.org

14. “You get pretty good at throwing starfish into deeper water.” asnailodyessy.com Nature, 2010 California Academy of Sciences “I’ve always thought of myself as a wader due to my size.”

15. Kick-It-And-See Ecology • Changed ecology from an observational to an experimental science • 3 years on Makah Bay • 8 x 2 m plots • Removal of Pisasterochraceus • Unmanipulated control • Transect lines to measure density of resident macroinvertebrates and benthic algae scientificamerican.com Bruno, 2007 ppt

16. What Maintains Diversity? • Previously, thought diversity = ecosystem stability • “Stability increases as the number of links increase” (MacArthur, 1955) • “A rich fauna and flora…tends to be very stable because of multiplicity of ecological checks and balances” (Watt, 1964) • Paine – absence of one individual can shift entire population into monoculture • Species richness decreased from 15 to 8 online.santarosa.edu

17. Paine, 1966 Nature Education, 2010

18. 3 Pages That Changed the World (of Ecology) • Loss can initiate trophic cascades: the rise and fall of connected species throughout the food web

19. Type of Keystones • Predators – Sea Otters, Gray Wolves • Prey • Mutualists – hummingbirds • Hosts – Saguaro cactus • Parasites • Modifiers – N. American Beaver • Pollinators • Where does it end?

20. Challenges include context dependency and diversity dictating keystone status • 1994 – United Nations’ Global Biodiversity Assessment met to identify issues and challenges with keystone species concept • 9 “keystone cops” including Paine, Tilman, Mary Power, Bruce Menge • Written as discussion between “Dr. Knowitall,” “Empiricist” and “Skeptic”

21. A keystone species is one that has a disproportionately large effect on its environment relative to its abundance, size or biomass Goodness gracious, it’s Pisasterochraceus

22. Drawing from 1999 • Paul Dayton • Jane Lubchenco • Bruce Menge • Steve Palumbi Marian Kohn, 1999 (Nature, 2013)

25. Paine = a Keystone The New York Times, 2012

26. Questions? sfbbo.org

27. References Mills, L. Scott, Michael E. Soule, and Daniel F. Doak. "The keystone-species concept in ecology and conservation." BioScience 43.4 (1993): 219-224. Levin, Simon A., and Robert T. Paine. "Disturbance, patch formation, and community structure." Proceedings of the National Academy of Sciences 71.7 (1974): 2744-2747. Paine, Robert T. "Food web complexity and species diversity." American Naturalist (1966): 65-75. Paine, Robert T. "A note on trophic complexity and community stability." The American Naturalist 103.929 (1969): 91-93. Paine, Robert T. "Food webs: linkage, interaction strength and community infrastructure." Journal of Animal Ecology 49.3 (1980): 667-685. Paine, Robert T. "A conversation on refining the concept of keystone species."Conservation Biology 9.4 (1995): 962- 964. Paine, Robert T., et al. "Trouble on oiled waters: lessons from the Exxon Valdez oil spill." Annual Review of Ecology and Systematics (1996): 197-235. Power, Mary E., et al. "Challenges in the quest for keystones." BioScience46.8 (1996): 609-620.