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“All Animals Weren’t Created Equal”
Robert T. Paine
Jenessa Kay
April 23, 2013
Community Ecology
Nature Education, 2012
Biographical Info
 Born in Cambridge, MA – April 13,1933
 Happy belated birthday!
 AB - Harvard University, 1954
 Army Battalion Gardener
Kinne (ed.), 1994
 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
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
Cited by 776
Cited by 3652
Cited by 1178
Cited by 920
BioScience, Vol. 46, No. 8 (Sep., 1996)
Cited by 1017
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??
The New York Times, 2012
Kevin Schafer
Study Location
 Makah Bay – mainland WA
 Tatoosh Island - 0.5 miles
offshore from Cape Flattery
 Part of Makah Reservation
makah.com
 Longest ongoing study of a
single area by the same scientist
in the U.S.
Superstock.com
Tatoosh Island
legendsofamerica.com
wikimedia.com
history.noaa.gov (1943)
“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.”
The New York Times, 2012
“His intellectual
presence is so
commanding that his
physical presence
hardly registers.”
naturalhistoriesproject.org
“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
“What would happen if
we removed the top
predator from an
ecosystem?”
naturalhistoriesproject.org
“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.”
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 Pisaster ochraceus
 Unmanipulated control
 Transect lines to measure density
of resident macroinvertebrates
and benthic algae
scientificamerican.com
Bruno, 2007 ppt
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
Paine, 1966
Nature Education, 2010
3 Pages That Changed the World (of Ecology)
 Loss can initiate trophic
cascades: the rise and fall of
connected species throughout
the food web
Type of Keystones
 Predators – Sea Otters, Gray Wolves
 Prey
 Mutualists – hummingbirds
 Hosts – Saguaro cactus
 Parasites
 Modifiers – N. American Beaver
 Pollinators
 Where does it end?
 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”
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 Pisaster ochraceus
 Drawing from 1999
 Paul Dayton
 Jane Lubchenco
 Bruce Menge
 Steve Palumbi
Marian Kohn, 1999 (Nature, 2013)
Paine = a Keystone
The New York Times, 2012
Questions?
sfbbo.org
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): 962964.
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.