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By: Stephen C. Wagner (Department of Biology, Stephen F. Austin State University) 2010 Nature Education
Citation: Wagner, S. C. (2010) Keystone Species. Nature Education Knowledge 3(10):51
A keystone in an arch's crown secures the other stones in place. Keystone species play
the same role in many ecological communities by maintaining the structure and integrity
of the community.
Aa Aa Aa
Paine's Milestones
The term keystone species was first coined by Robert Paine (1966) after extensive studies examining the interaction
strengths of food webs in rocky intertidal ecosystems in the Pacific Northwest. One of his study sites, located at Mukkaw
Bay, contained a community consistently dominated by the same species of mussels, barnacles, and the starfish, Pisaster
ochraceus, which preys upon the other species as a top predator (Figure 1).
Paine (1966) had observed that the diversity of organisms in rocky intertidal ecosystems declined as the number of
predators in those ecosystems decreased. He hypothesized that some of these consumers might be playing a greater role
than others in controlling the numbers of species coexisting in these communities. He tested his hypothesis in an experiment
that involved selecting a "typical" piece of shoreline at Mukkaw Bay, about 8 meters long by 2 meters wide, that was kept
free of starfish. This area was compared to an adjacent, undisturbed control area of equal size.
Figure 2: Effect of removal of top predator on total species number in intertidal ecosystem
2010 Nature Education All rights reserved.
In his seminal paper that followed this work, Paine (1969) derived the term keystone species to describe the starfish in these
intertidal ecosystems. Of these species he commented: "The species composition and physical appearance were greatly
modified by the activities of a single native species high in the food web. These individual populations are the keystone of
the community's structure, and the integrity of the community and its unaltered persistence through time."
Paine went on to describe the criteria for a keystone species. A keystone species exerts top-down influence on lower trophic
levels and prevents species at lower trophic levels from monopolizing critical resources, such as competition for space or
key producer food sources. This paper represented a watershed in the description of ecological relationships between
species. In the twenty years that followed its publication, it was cited in over ninety publications. Additionally, the original
paper describing the intertidal areas was cited in over 850 papers during the same time period (Mills et al. 1993).
hosts because they provide habitat for a variety of other species. Keystone prey are species that can maintain their numbers
despite being preyed upon, therefore controlling the density of a predator.
Figure 4: The gray wolf (Canis lupus), a keystone species in Yellowstone National Park
2010 Nature Education All rights reserved.
When the federal government set aside the Greater Yellowstone Ecosystem (GYE) as a national park in 1872, about three to
four hundred wolves were present, preying mostly upon large hooved ungulates such as elk (Cervus canadensis, Figure 5)
and bison (Yellowstone Association 1996). Fearing the wolves' impact on elk and bison herds as well as livestock owned by
area ranchers, the federal government began eradicating the wolf population. Bounty programs that continued until 1965
offered as much as $50 per wolf. Wolves were trapped, shot, dug from their dens, hunted with dogs, and poisoned. In
Yellowstone National Park, park rangers killed the last two remaining pups in 1924. By the 1930s wolves had been
effectively eliminated from the contiguous 48 States and Mexico and only remained in high numbers in Alaska.
Figure 5: The elk (Cervus canadensis), principal prey of gray wolves (Canis lupus)
2010 Nature Education All rights reserved.
With their primary predator eliminated, elk populations exploded, leading to the overgrazing of plants, especially those found
in riparian zones (Laliberte & Ripple 2004). Significant declines in the populations of many plant species (e.g., aspen, willow)
resulted, which in turn influenced other wildlife, such as beaver and songbird populations (Ripple & Breschetta 2004,
Hallofsky & Ripple 2008). Intensive browsing of aspen (Populus tremuloides) stands, for example, led to a rapid decline in
the number of seedlings and root sprouts growing into saplings and trees. For many stands of these trees, only large
diameter trees (i.e., those that had matured before the wolves were eradicated) remained.
Disappearance of these and other plant species not only caused the loss of habitat for many other animals but also
influenced other ecological factors (Smith et al. 2009), including stream bank stability, the deposition of organic matter and
fine sediment in riparian zones, water temperature regulation via shading, and nutrient cycling. The removal of wolves thus
led to the instability of riparian and other environmentally sensitive areas.
Summary
The concept of keystone species was first proposed and demonstrated in the 1960s by the dominance of top-predator
starfish in intertidal ecosystems. Keystone species are species that play a disproportionately large role in the prevalence and
population levels of other species within their ecosystem or community. The recovery of the gray wolf after its eradication
from Yellowstone National Park, almost ninety years ago, demonstrates how crucial keystone species are to the long-term
sustainability of the ecosystems they inhabit. Most importantly, the preservation and restoration of keystone species is
essential for maintaining and/or reestablishing the historic structure and function of the ecosystems they inhabit.
Not all species are equal with respect to their [effects on other species. Starfish feeding in the intertidal zone
clean an area free of barnacles and mussels. These barnacles and mussels, without predation by the
starfish, would come to dominate the community. In a classic 1966 study, Robert Paine removed starfish
from enclosures. In those enclosures where the starfish were removed, the number of species in the
community dropped from fifteen to eight. Paine called starfish a keystone species, one whose presence
has a dramatic effect on species diversity.
Prior to 1973 most ecologists thought that more diverse ecosystems would be more stable than would ones
with fewer species. This general belief, what has become known as the diversity-stability hypothesis,
was based on a variety of observations but not really tested. One such observation was that cultivated land
that had simplified ecological communities was more subject to species invasions than similar areas that
hadn't had human influence. In addition, insect outbreaks are much more common in the less diverse boreal
forests than they are in tropical forests.
In 1973 Robert May published a theoretical study that challenged the intuitive ideas that ecologists had
about the diversity-stability hypothesis. May analyzed randomly constructed communities and found that
communities with more species tended to be less, not more, stable. May's study, like more theoretical
studies of the 1970s, assumed that population numbers of each species were at equilibria. This assumption
was made, not because it reflected reality, but because it made the mathematics more tractable. More
recent studies have shown that if there is some degree of flux in the population numbers, the community
can maintain more species than in equilibrium. This variability may allow different species to respond
differently to the environment, and can result in fewer species being lost due to competitive exclusion. When
theoretical ecologists relax the equilibrium assumption and allow for population fluxes, they have found
results consistent with the diversity-stability hypothesis: communities with more species are more stable.
Several lines of evidence now support the diversity-stability hypothesis. The studies conducted by David
Tilman and his colleagues provide some of the strongest evidence for the hypothesis. In 1982 Tilman divided
grassland fields in Minnesota's Cedar Creek National History Area into more than 200 plots. He and his
colleagues monitored the species richness and community biomass (the total mass of all plants) in each of
those fields over the next two decades. They found that diversity within a community is positively correlated
with plant community stability, as defined by the extent of variation in community biomass. Various other
studies at different scales have found similar results: stability increases with diversity.4
kelp beds and associated marine life there. That's because one of the favorite
delicacies of the otter is the large sea urchin, which in turn feeds on kelp. As the sea
otter returns to its native territories, scientists expect the population of invertebrates,
like urchins and abalone, for example, to decrease as marine plant biomass increases.
In fact, a decrease in sport and commercial abalone fisheries was reported following
an influx of sea otters into areas of California, causing a controversy there.
Paine's work has been cited by hundreds of researchers over the years, and it has been
proposed as a foundation for management efforts to protect the biological diversity of
the world's ecosystems.
"Its importance," says Paine, "is that it convinced managers and conservationists alike
that the ecological impact of single species matters. That is, in order to manage,
understand, and restore ecological assemblages, the roles of individual species have to
be understood and considered."
The concept has provided a powerful model for understanding the forces that organize
ecological communities, and it has influenced the thinking of managers and policy
makers as they set priorities in their efforts to conserve species and habitats. Over the
years, controversy in scientific circles has grown up around the term keystone species
as a result of "overly expansive usage." Through a United Nations Environmental
Programme workshop held in 1994, prominent researchers in the field reached a new
consensus about a definition of the term and new guidelines for its expanding
applications in the future.
1.
"The Keystone-Species Concept in Ecology and Conservation," L. Scott Mills, Michael E. Soule, and
Daniel F. Doak, BioScience, 43 (4), 219 (1993).
2.
"Sea Otters: Their Role in Structuring Nearshore Communities," Science, 185, 1058 (1974).
3.
"The Keystone Cops Meet in Hilo," Mary E. Power and L. Scott Mills, TREE, 10 (5), 182 (1995).