Keystone species

A keystone species is a species that has a disproportionate effect on its environment relative to its abundancecite journal
author=Paine, R.T.
year=1995
title=A Conversation on Refining the Concept of Keystone Species
journal=Conservation Biology
volume=9
issue=4
pages=962–964
doi=10.1046/j.1523-1739.1995.09040962.x] . Such species affect many other organisms in an ecosystem and help to determine the types and numbers of various others species in a community.

Such an organism plays a role in its ecosystem that is analogous to the role of a keystone in an arch. While the keystone feels the least pressure of any of the stones in an arch, the arch still collapses without it. Similarly, an ecosystem may experience a dramatic shift if a keystone species is removed, even though that species was a small part of the ecosystem by measures of biomass or productivity. It has become a very popular concept in conservation biology.cite journal
author=Mills, L.S.
coauthors=Soule, M.E.; Doak, D.F.
year=1993
title=The Keystone-Species Concept in Ecology and Conservation
journal=BioScience
volume=43
issue=4
pages=219–224
url=http://links.jstor.org/sici?sici=0006-3568(199304)43%3A4%3C219%3ATKCIEA%3E2.0.CO%3B2-E
accessdate=2007-10-04
doi=10.2307/1312122]

Definition

The definition given here is somewhat qualitative in nature because there is not yet an accepted, rigorous definition.cite journal
author=Power, M.E.
coauthors=Tilman, D.; Estes, J.A.; Menge, B.A.; Bond, W.J.; Mills, L.S.; Daily, G.; Castilla, J.C.; Lubchenco, J.; Paine, R.T.
year=1996
title=Challenges in the Quest for Keystones
journal=BioScience
volume=46
issue=8
pages=609–620
url=http://links.jstor.org/sici?sici=0006-3568(199609)46%3A8%3C609%3ACITQFK%3E2.0.CO%3B2-7
accessdate = 2007-10-04
doi=10.2307/1312990] An issue is how to measure both abundance and impact and at what point to draw the line. Abundance in this context can be measured by biomass or productivity, among other metrics. Impact is even harder to define. It has been suggested that such keystone predators can be identified in ecosystems by their biomass dominance within ecological functional groups, even though they may be relatively rare in relation to the ecosystem as a whole.cite journal
author=Davic, R.D.
year=2003
title=Linking Keystone Species and Functional Groups: A New Operational Definition of the Keystone Species Concept
journal=Conservation Ecology
volume=7
issue=1
pages=r11
url=http://www.ecologyandsociety.org/vol7/iss1/resp11/
accessdate=2007-10-04]

The term is often misused in places where foundation species would be more appropriate. A keystone species is not simply one whose disappearance would alter the ecosystem. Removing any abundant species from any ecosystem will drastically alter that ecosystem by definition. What makes the concept of keystone species attractive to conservationists is that a species with a relatively tiny physical footprint can be critical to the health of its ecosystem.

Examples

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Without a consensus on its exact definition, we are left to illustrate the concept of keystone species with a list of examples.

A classic keystone species is a small predator that prevents a particular herbivorous species from eliminating dominant plant species. Since the prey numbers are low, the keystone predator numbers can be even lower and still be effective. Yet without the predators, the herbivorous prey would explode in numbers, wipe out the dominant plants, and dramatically alter the character of the ecosystem. The exact scenario changes in each example, but the central idea remains that through a chain of interactions, a non-abundant species has an out-sized impact on ecosystem functions. One example is the weevil and its suggested keystone effects on aquatic plant species diversity by prey activities on nuisance Euransian Watermilfoil.cite journal
author=Creed Jr, R.P.
year=2000
title=Is there a new keystone species in North American lakes and rivers?
journal=OIKOS
volume=91
issue=2
doi=10.1034/j.1600-0706.2000.910222.x
pages=405]

Predators

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Some starfish may perform this function by preying on sea urchins, mussels, and other shellfish that have no other natural predators. If the sea star is removed from the ecosystem, the mussel population explodes uncontrollably, driving out most other species, while the urchin population annihilates coral reefs. In his classic 1966 paper, Dr. Robert Paine described such a system in Mukkaw Bay in Washington State.cite journal
author=Paine, R.T.
year=1966
title=Food Web Complexity and Species Diversity
journal=The American Naturalist
volume=100
issue=910
pages=65–75
url=http://links.jstor.org/sici?sici=0003-0147(196601/02)100:910%3C65:FWCASD%3E2.0.CO;2-D
accessdate=2007-10-04
doi=10.1086/282400] This led to his 1969 paper where he proposed the keystone species concept.cite journal
author=Paine, R.T.
year=1969
title=A Note on Trophic Complexity and Community Stability
journal=The American Naturalist
volume=103
issue=929
pages=91–93
url=http://links.jstor.org/sici?sici=0003-0147(196901%2F02)103%3A929%3C91%3AANOTCA%3E2.0.CO%3B2-U
accessdate = 2007-10-04
doi=10.1086/282586]

Similarly, sea otters in kelp forests keep sea urchins in check. Kelp roots are merely anchors, and not the vast nutrient gathering networks of land plants. Thus the urchins only need to eat the roots of the kelp, a tiny fraction of the plant's biomass, to remove it from the ecosystem.cite journal
last=Estes
first=James E.
coauthors=Norman S. Smith, John F. Palmisano
year=1978
title=Sea otter predation and community organization in the Western Aleutian Islands, Alaska
journal=Ecology
volume=59
issue=4
pages=822–833
url=http://links.jstor.org/sici?sici=0012-9658(197822)59%3A4%3C822%3ASOPACO%3E2.0.CO%3B2-N
accessdate=2007-10-04
doi=10.2307/1938786] cite journal
author=Cohn, J.P.
year=1998
title=Understanding Sea Otters
journal=BioScience
volume=48
issue=3
pages=151–155
url=http://links.jstor.org/sici?sici=0006-3568(199803)48%3A3%3C151%3AUSO%3E2.0.CO%3B2-G
accessdate=2007-10-04
doi=10.2307/1313259]

These creatures need not be apex predators. Sea stars are prey for sharks, rays, and sea anemones. Sea otters are prey for orca.cite journal
author=Estes, J.A.
coauthors=Tinker, M.T.; Williams, T.M.; Doak, D.F.
date=1998-10-16
title=Killer whale predation on sea otters linking oceanic and nearshore ecosystems
journal=Science
volume=282
issue=5388
pages=473–476
doi=10.1126/science.282.5388.473
url=http://adsabs.harvard.edu/abs/1998Sci...282..473E
accessdate = 2007-10-04
pmid=9774274]

Engineers

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In North America, the grizzly bear is a keystone species - not as a predator but as ecosystem engineers. They transfer nutrients from the oceanic ecosystem to the forest ecosystem. The first stage of the transfer is performed by salmon, rich in nitrogen and potassium, who swim up rivers, sometimes for hundreds of miles. The bears then capture the salmon and carry them onto dry land, dispersing nutrient-rich feces and partially-eaten carcasses. It has been estimated that the bears leave up to half of the salmon they harvest on the forest floor.Fact|date=October 2007

Another ecosystem engineering keystone species is the beaver, which transforms its territory from a stream to a pond or swamp.cite journal
author=Wright, J.P.
coauthors=Jones, C.G.; Flecker, A.S.
year=2002
title=An ecosystem engineer, the beaver, increases species richness at the landscape scale
journal=Oecologia
volume=132
issue=1
pages=96–101
url=http://www.springerlink.com/index/0637GF0979LRU90J.pdf
accessdate = 2007-10-04
doi=10.1007/s00442-002-0929-1]

In the African savanna, the larger herbivores, especially the elephants, shape their environment. The elephants destroy trees, making room for the grass species. Without these animals, much of the savanna would turn into woodland.cite book
last=Leakey
first=Richard
authorlink=Richard Leakey
coauthors=Roger Lewin
title=The sixth extinction: biodiversity and its survival
origdate=1995
accessdate=2007-10-28
year=1999
publisher=Phoenix
location=London
isbn=1-85799-473-6
pages=pp. 216-217
chapter=11 The modern elephant story]

References

ee also

*Foundation species
*Indicator species
*Flagship species
*Indigenous
*Introduced species

External links

* [http://www.washington.edu/research/pathbreakers/1969g.html A summary of Paine's original hypothesis from the University of Washington's Pathbreaker series]